Preformed aerenchyma determines the differential tolerance response under partial submergence imposed by fresh and saline water flooding in rice.

Plant's response to fresh- and saline-water flooding and the resulting partial submergence, seems different due to the added complexities of element toxicity of salinity. We identified a few rice genotypes which can tolerate combined stresses of partial submergence and salinity during saline water flooding. To gain mechanistic insights, we compared two rice genotypes: Varshadhan (freshwater-flooding tolerant) and Rashpanjor (both fresh- and saline-water flooding tolerant). We found greater ethylene production and increased "respiratory burst oxidase homolog" (RBOH)-mediated reactive oxygen species (ROS) production led to well-developed constitutive aerenchyma formation in Rashpanjor, which makes it preadapted to withstand fresh- and saline-water flooding. On the contrary, an induced aerenchyma formation-dependent tolerance mechanism of Varshadhan worked well for freshwater flooding but failed to provide tolerance to saline-water flooding. Additional salt stress was found to significantly inhibit the induced aerenchyma formation process due to the dampening of ROS signaling by the action of metallothionein in Varshadhan. Besides, inconspicuous changes in ionic regulation processes in these two genotypes under saline-water flooding suggest preadapted constitutive aerenchyma formation plays a more significant role than elemental toxicity per se in tolerating combined stresses encountered during saline water flooding in rice. Overall, our study indicated that well-developed constitutive aerenchyma provide an adaptive advantage during partial submergence due to saline water flooding in rice as the key process of induced aerenchyma formation is hampered in the presence of salinity stress coupled with partial submergence.

Recent Advances of Genetic Resources, Genes and Genetic Approaches for Flooding Tolerance in Rice.

Flooding is one of the most hazardous natural disasters and a major stress constraint to rice production throughout the world, which results in huge economic losses. The frequency and duration of flooding is predicted to increase in near future as a result of global climate change. Breeding of flooding tolerance in rice is a challenging task because of the complexity of the component traits, screening technique, environmental factors and genetic interactions. A great progress has been made during last two decades to find out the flooding tolerance mechanism in rice. An important breakthrough in submergence research was achieved by the identification of major quantitative trait locus (QTL) SUB1 in rice chromosomes that acts as the primary contributor for tolerance. This enabled the use of marker-assisted backcrossing (MABC) to transfer SUB1 QTL into popular varieties which showed yield advantages in flood prone areas. However, SUB1 varieties are not always tolerant to stagnant flooding and flooding during germination stage. So, gene pyramiding approach can be used by combining several important traits to develop new breeding rice lines that confer tolerances to different types of flooding. This review highlights the important germplasm/genetic resources of rice to different types of flooding stress. A brief discussion on the genes and genetic mechanism in rice exhibited to different types of flooding tolerance was discussed for the development of flood tolerant rice variety. Further research on developing multiple stresses tolerant rice can be achieved by combining SUB1 with other tolerance traits/genes for wider adaptation in the rain-fed rice ecosystems.

Rice with SUB1 QTL possesses greater initial leaf gas film thickness leading to delayed perception of submergence stress.

BACKGROUND AND AIMS: Submergence tolerance in rice is primarily attributed to the action of the SUB1 gene, but other associated traits such as leaf gas film (LGF) thickness, leaf hydrophobicity, porosity and leaf density have been known to aid submergence tolerance in rice. However, association of these traits with SUB1 quantitative trait locus (QTL) has not been demonstrated. In this study, we aim to investigate (1) whether the presence of the SUB1 QTL in the genetic background has any influence on the thickness of the LGF and (ii) whether its removal has any impact on stress perception and submergence tolerance in Sub1 and non-Sub1 rice. METHODS: We examined 12 genotypes (including both Sub1 and non-Sub1 types) for different leaf traits such as initial LGF thickness, leaf hydrophobicity, tissue porosity and leaf density in order to work out the relatioship of these traits to the SUB1 QTL in rice. Furthermore, we investigated the changes in the gene expression profile and different metabolic processes in selected genotypes in the presence and absence of their LGF to study its impact on stress perception and adaptation. KEY RESULTS: The initial thickness of the LGF and hydrophobicity seemed to have a highly positive correlation with the presence of the SUB1 QTL in the genetic background of rice; however, other leaf traits such as porosity and density seemed to be independent of it. Artificial removal of the LGF resulted in partial loss of tolerance, showing increased ethylene production and early induction of anoxia-related genes (SUB1A-1, ACS5, Ramy3D and ADH1) which manifested symptoms such as increased stem elongation, faster chlorophyll and starch breakdown, and partial loss of quiescence in SUB1-containing rice genotypes. Stripping of the LGF resulted in early and enhanced induction of SUB1A-1, indicating a quicker perception of stress. CONCLUSIONS: The presence of SUB1 in the genetic background positively influences surface hydrophobicity and the concomitant LGF thickness of rice. Furthermore, LGF helps in terms of providing better ethylene dissipation and reduced in planta accumulation, owing to the slowing down of ethylene-induced leaf senescence under submergence stress.

Tissue Tolerance Coupled With Ionic Discrimination Can Potentially Minimize the Energy Cost of Salinity Tolerance in Rice.

Salinity is one of the major constraints in rice production. To date, development of salt-tolerant rice cultivar is primarily focused on salt-exclusion strategies, which incur greater energy cost. The present study aimed to evaluate a balancing strategy of ionic discrimination vis-à-vis tissue tolerance, which could potentially minimize the energy cost of salt tolerance in rice. Four rice genotypes, viz., FL478, IR29, Kamini, and AC847, were grown hydroponically and subjected to salt stress equivalent to 12 dS m-1 at early vegetative stage. Different physiological observations (leaf chlorophyll content, chlorophyll fluorescence traits, and tissue Na+ and K+ content) and visual scoring suggested a superior Na+-partitioning strategy operating in FL478. A very low tissue Na+/K+ ratio in the leaves of FL478 after 7 days of stress hinted the existence of selective ion transport mechanism in this genotype. On the contrary, Kamini, an equally salt-tolerant genotype, was found to possess a higher leaf Na+/K+ ratio than does FL478 under similar stress condition. Salt-induced expression of different Na+ and K+ transporters indicated significant upregulation of SOS, HKT, NHX, and HAK groups of transporters in both leaves and roots of FL478, followed by Kamini. The expression of plasma membrane and vacuolar H+ pumps (OsAHA1, OsAHA7, and OsV-ATPase) were also upregulated in these two genotypes. On the other hand, IR29 and AC847 showed greater salt susceptibility owing to excess upward transport of Na+ and eventually died within a few days of stress imposition. But in the "leaf clip" assay, it was found that both IR29 and Kamini had high tissue-tolerance and chlorophyll-retention abilities. On the contrary, FL478, although having higher ionic-discrimination ability, showed the least degree of tissue tolerance as evident from the LC50 score (amount of Na+ required to reduce the initial chlorophyll content to half) of 336 mmol g-1 as against 459 and 424 mmol g-1 for IR29 and Kamini, respectively. Overall, the present study indicated that two components (ionic selectivity and tissue tolerance) of salt tolerance mechanism are distinct in rice. Unique genotypes like Kamini could effectively balance both of these strategies to achieve considerable salt tolerance, perhaps with lesser energy cost.

Ionic selectivity and coordinated transport of Na+ and K+ in flag leaves render differential salt tolerance in rice at the reproductive stage.

MAIN CONCLUSION: The present study shows that salt tolerance in the reproductive stage of rice is primarily governed by the selective Na+ and K+ transport from the root to upper plant parts. Ionic discrimination at the flag leaf, governed by differential expression of Na+- and K+-specific transporters/ion pumps, is associated with reduced spikelet sterility and reproductive stage salt tolerance. Reproductive stage salt tolerance is crucial in rice to guarantee yield under saline condition. In the present study, differential ionic selectivity and the coordinated transport (from root to flag leaf) of Na+ and K+ were investigated to assess their impact on reproductive stage salt tolerance. Four rice genotypes having differential salt sensitivity were subjected to reproductive stage salinity stress in pots. The selective Na+ and K+ transport from the root to upper plant parts was observed in tolerant genotypes. We noticed that prolonged salt exposure did not alter flag leaf greenness even up to 6 weeks; however, it had a detrimental effect on panicle development especially in the salt-susceptible genotype Sabita. But more precise chlorophyll fluorescence imaging analysis revealed salinity-induced damages in Sabita. The salt-tolerant genotype Pokkali (AC41585), a potential Na+ excluder, managed to sequester higher Na+ load in the roots with little upward transport as evident from greater expression of HKT1 and HKT2 transporters. In contrast, the moderately salt-tolerant Lunidhan was less selective in Na+ transport, but possessed a higher capacity to Na+ sequestration in leaves. Higher K+ uptake and tissue-specific redistribution mediated by HAK and AKT transporters showed robust control in selective K+ movement from the root to flag leaf and developing panicles. On the contrary, expressions of Na+-specific transporters in developing panicles were either down-regulated or unaffected in tolerant and moderately tolerant genotypes. Yet, in the panicles of the susceptible genotype Sabita, some of the Na+-specific transporter genes (SOS1, HKT1;5, HKT2;4) were upregulated. Apart from the ionic regulation strategy, cellular energy balance mediated by different plasma-membrane and tonoplastic H+-pumps were also associated with the reproductive stage salt tolerance in rice.

Riboflavin (Vitamin B2) mediated defence induction against bacterial leaf blight: probing through chlorophyll a fluorescence induction O-J-I-P transients.

Bacterial leaf blight (BLB) is a serious threat for rice (Oryza sativa L.) cultivation caused by the bacterial pathogen Xanthomonas oryzae pv. oryzae. The pathogen mainly damages the leaf chlorophyllous tissue, resulting in poor photosynthesis and causing up to 50% reductions in grain yield. In the present work, we have compared the structural and functional ability of the chloroplast of three varieties of rice with different degrees of susceptibility (TN1, highly susceptible; IR-20, moderately resistant; DV-85, resistant to BLB) treated with riboflavin (1 and 2mM) and infected with BLB, with chlorophyll fluorescence as a tool. As indicated by the chlorophyll fluorescence technique, the disease progress curve and yield data, riboflavin acted as an effective vitamin for inducing resistance against BLB. Plants treated with riboflavin showed improved PSII activity, more chlorophyll content and higher yield than the diseased plants.