Novel QTL for Lateral Root Density and Length Improve Phosphorus Uptake in Rice (Oryza sativa L.).

The rice root system consists of two types of lateral roots, indeterminate larger L-types capable of further branching, and determinate, short, unbranched S-types. L-type laterals correspond to the typical lateral roots of cereals whereas S-type laterals are unique to rice. Both types contribute to nutrient and water uptake and genotypic variation for density and length of these laterals could be exploited in rice improvement to enhance adaptations to nutrient and water-limited environments. Our objectives were to determine how best to screen for lateral root density and length and to identify markers linked to genotypic variation for these traits. Using different growing media showed that screening in nutrient solution exposed genotypic variation for S-type and L-type density, but only the lateral roots of soil-grown plants varied for their lengths. A QTL mapping population developed from parents contrasting for lateral root traits was grown in a low-P field, roots were sampled, scanned and density and length of lateral roots measured. One QTL each was detected for L-type density (LDC), S-type density on crown root (SDC), S-type density on L-type (SDL), S-type length on L-type (SLL), and crown root number (RNO). The QTL for LDC on chromosome 5 had a major effect, accounting for 46% of the phenotypic variation. This strong positive effect was confirmed in additional field experiments, showing that lines with the donor parent allele at qLDC5 had 50% higher LDC. Investigating the contribution of lateral root traits to P uptake using stepwise regressions indicated LDC and RNO were most influential, followed by SDL. Simulating effects of root trait differences conferred by the main QTL in a P uptake model confirmed that qLDC5 was most effective in improving P uptake followed by qRNO9 for RNO and qSDL9 for S-type lateral density on L-type laterals. Pyramiding qLDC5 with qRNO9 and qSDL9 would be possible given that trade-offs between traits were not detected. Phenotypic selection for the RNO trait during variety development would be feasible, however, the costs of doing so reliably for lateral root density traits is prohibitive and markers identified here therefore provide the first opportunity to incorporate such traits into a breeding program.

Corrigendum: QTL mapping for early root and shoot vigor of upland rice (Oryza sativa L.) under P deficient field conditions in Japan and Madagascar.

[This corrects the article DOI: 10.3389/fpls.2022.1017419.].

Mechanisms of ozone tolerance in rice: characterization of two QTLs affecting leaf bronzing by gene expression profiling and biochemical analyses.

High surface ozone concentration is increasingly being recognized as a factor that negatively affects crop yields in Asia. However, little progress has been made in developing ozone-tolerant genotypes of rice-Asia's major staple crop. This study aimed to identify possible tolerance mechanisms by characterizing two quantitative trait loci (QTLs) that were previously shown to influence visible leaf symptoms under ozone exposure (120 nl l(-1), 7 h d(-1), 13 d). Two chromosome segment substitution lines (SL15 and SL41) that carried introgressions of the QTLs OzT3 and OzT9, respectively, were exposed to ozone at 120 nl l(-1) along with their parent Nipponbare. In accordance with the expected QTL effect, SL15 showed stronger visible symptoms of ozone damage than Nipponbare, whereas SL41 had fewer symptoms. Gene expression profiling by microarray hybridization yielded 470 probes that were differentially expressed in SL15 and 314 in SL41. Potential tolerance mechanisms were evaluated by investigating changes in gene expression in three general categories. (i) Processes involved in programmed cell death, in which a number of genes related to ethylene or jasmonic acid metabolism or general disease resistance were identified that were differentially regulated in one of the substitution lines. (ii) Biosynthesis of antioxidants. Testing this hypothesis did not reveal any genes differentially regulated between genotypes, and it was thus rejected. (iii) Turnover of antioxidants and enzymatic detoxification of radical oxygen species (ROS), in which a number of differentially regulated genes were also identified. Genes encoding antioxidant enzymes (catalase and peroxidases) tended to be more strongly expressed in SL15. A potential tolerance gene which encodes a putative ascorbate oxidase was identified within the QTL introgression in SL41. This gene showed consistently lower expression in SL41 under ozone exposure across different points in time within independent experiments. Its expression may be involved in mechanisms leading to enhanced ascorbic acid status in SL41 under ozone exposure, and may be linked to a higher concentration of total apoplastic ascorbic acid in SL41 that was observed in an independent experiment.

Comparative sequence analyses of the major quantitative trait locus phosphorus uptake 1 (Pup1) reveal a complex genetic structure.

The phosphorus uptake 1 (Pup1) locus was identified as a major quantitative trait locus (QTL) for tolerance of phosphorus deficiency in rice. Near-isogenic lines with the Pup1 region from tolerant donor parent Kasalath typically show threefold higher phosphorus uptake and grain yield in phosphorus-deficient field trials than the intolerant parent Nipponbare. In this study, we report the fine mapping of the Pup1 locus to the long arm of chromosome 12 (15.31-15.47 Mb). Genes in the region were initially identified on the basis of the Nipponbare reference genome, but did not reveal any obvious candidate genes related to phosphorus uptake. Kasalath BAC clones were therefore sequenced and revealed a 278-kbp sequence significantly different from the syntenic regions in Nipponbare (145 kb) and in the indica reference genome of 93-11 (742 kbp). Size differences are caused by large insertions or deletions (INDELs), and an exceptionally large number of retrotransposon and transposon-related elements (TEs) present in all three sequences (45%-54%). About 46 kb of the Kasalath sequence did not align with the entire Nipponbare genome, and only three Nipponbare genes (fatty acid alpha-dioxygenase, dirigent protein and aspartic proteinase) are highly conserved in Kasalath. Two Nipponbare genes (expressed proteins) might have evolved by at least three TE integrations in an ancestor gene that is still present in Kasalath. Several predicted Kasalath genes are novel or unknown genes that are mainly located within INDEL regions. Our results highlight the importance of sequencing QTL regions in the respective donor parent, as important genes might not be present in the current reference genomes.

Genotypic variation in tolerance to elevated ozone in rice: dissection of distinct genetic factors linked to tolerance mechanisms.

Tropospheric ozone concentrations are increasing in many Asian countries and are expected to reach levels that adversely affect crop production. Developing ozone-tolerant rice (Oryza sativa L.) varieties is therefore essential to prevent yield losses in the future. The aims of this study were to assess genotypic variation for ozone tolerance in rice, to identify quantitative trait loci (QTL) conferring tolerance, and to relate QTLs to physiological tolerance mechanisms. The response of 23 varieties to elevated ozone (120 nl l(-1)) was assessed based on leaf bronzing and dry weight loss. The traditional variety 'Kasalath' was highly tolerant, whereas the modern variety 'Nipponbare' showed significant dry weight reductions. Using the Nipponbare/Kasalath/Nipponbare mapping population, six QTLs associated with tolerance to elevated ozone were identified, of which three were subsequently confirmed in Nipponbare/Kasalath substitution lines (SLs). Two QTLs associated with leaf bronzing were located on chromosomes three and nine. Kasalath alleles on chromosome three increased bronzing, while alleles on chromosome nine reduced bronzing. SLs carrying these contrasting QTLs differed significantly in leaf ascorbic acid (AsA) content when exposed to ozone, suggesting AsA as a principal antioxidant counteracting ozone-induced oxidative damage. A further confirmed QTL related to dry weight was located on chromosome eight, where the Kasalath allele increased relative dry weight. A SL carrying this QTL exhibited a less reduced net photosynthetic rate under ozone exposure compared with its recurrent parent Nipponbare. Although the effect of these QTLs on crop yield has not yet been established, their identification could be an important first step in developing ozone-tolerant rice varieties.