Tolerance of anaerobic conditions caused by flooding during germination and early growth in rice (Oryza sativa L.)

Abstract

Rice is semi-aquatic, adapted to a wide range of hydrologies, from aerobic soils in uplands to anaerobic and flooded fields in waterlogged lowlands, to even deeply submerged soils in flood-prone areas. Considerable diversity is present in native rice landraces selected by farmers over centuries. Our understanding of the adaptive features of these landraces to native ecosystems has improved considerably over the recent past. In some cases, major genes associated with tolerance have been cloned, such as SUB1A that confers tolerance of complete submergence and SNORKEL genes that control plant elongation to escape deepwater. Modern rice varieties are sensitive to flooding during germination and early growth, a problem commonly encountered in rainfed areas, but few landraces capable of germination under these conditions have recently been identified, enabling research into tolerance mechanisms. Major QTLs were also identified, and are being targeted for molecular breeding and for cloning. Nevertheless, limited progress has been made in identifying regulatory processes for traits that are unique to tolerant genotypes, including faster germination and coleoptile elongation, formation of roots and leaves under hypoxia, ability to catabolize starch into simple sugars for subsequent use in glycolysis and fermentative pathways to generate energy. Here we discuss the state of knowledge on the role of the PDC-ALDH-ACS bypass and the ALDH enzyme as the likely candidates effective in tolerant rice genotypes. Potential involvement of factors such as cytoplasmic pH regulation, phytohormones, reactive oxygen species scavenging and other metabolites is also discussed. Further characterization of contrasting genotypes would help in elucidating the genetic and biochemical regulatory and signaling mechanisms associated with tolerance. This could facilitate breeding rice varieties suitable for direct seeding systems and guide efforts for improving waterlogging tolerance in other crops.

Keywords: ALDH; alcoholic fermentation; anaerobic germination; direct seeding; flooding; hypoxia; pyruvate dehydrogenase bypass; submergence tolerance.

Figure 1

Left to right: seedlings of sensitive varieties “IR64” and “IR42” and tolerant landraces “Khao Hlan On” and “Khaiyan” 7 days after sowing in soil and flooding with 10 cm of water (from image collection of the International Rice Research Institute (IRRI) publicly available at Flickr http://www.flickr.com/photos/ricephotos/2199544108/in/photostream/).

Figure 2

Diagram summarizing carbohydrate metabolism under anoxic/hypoxic conditions in rice. Enzyme abbreviations: AlaAT, alanine aminotransferase (EC 2.6.1.2); LDH, lactate dehydrogenase (EC 1.1.1.27); PDC, pyruvate decarboxylase (EC 4.1.1.1); ADH, alcohol dehydrogenase (EC 1.1.1.1); mALDH, mitochondrial aldehyde dehydrogenase (EC 1.2.1.3); ACS, acetyl-CoA synthetase (EC 6.2.1.1); AR, ascorbate-free radical reductase (EC 1.6.5.4). ······indicates substrate movement and—indicates substrate conversion/modification. Reaction substrates are highlighted in bold letters.