Taiichiro Ookawa

New approach for rice improvement using a pleiotropic QTL gene for lodging resistance and yield

The use of fertilizer results in tall rice plants that are susceptible to lodging and results in reduced plant yields. In this study, using chromosome segment substitution lines, we identified an effective quantitative trait loci (QTL) for culm strength, which was designated STRONG CULM2 (SCM2). Positional cloning of the gene revealed that SCM2 was identical to ABERRANT PANICLE ORGANIZATION1 (APO1), a gene previously reported to control panicle structure. A near-isogenic line carrying SCM2 showed enhanced culm strength and increased spikelet number because of the pleiotropic effects of the gene. Although SCM2 is a gain-of-function mutant of APO1, it does not have the negative effects reported for APO1 overexpression mutants, such as decreased panicle number and abnormal spikelet morphology. The identification of lodging-resistance genes by QTL analysis combined with positional cloning is a useful approach for improving lodging resistance and overall productivity in rice.

The effects of irrigation regimes on the water use, dry matter production and physiological responses of paddy rice

In most cases, rice production is associated with flooding irrigation and the efficiency of irrigated water use (WUEi) is generally lower for production of rice than for other crops. We have examined the effects of various irrigation regimes on water consumption in a well-puddled paddy field, as well as on dry matter production, grain yield and physiological responses of the plants. Four sets of conditions were studied, with two replications, in the well-puddled paddy field: Continuous flooding irrigation treatment (CSF); three intermittent irrigation treatments, designated II-0, II-1 and II-2, in which plants were re-irrigated when the water potential of the soil fell below 0, –10, and –20 kPa at a depth of 5 cm, respectively. Water consumption was lower during II-0 than during CSF because the percolation rate was reduced by the reduction in the hydraulic head of the ponded water. Intermittent irrigation led to the repeated shrinking and swelling of soil during II-1 and II-2 and, therefore, soil cracks developed rapidly. Since they became the major routes of water percolation, these soil cracks increased water consumption during II-1 and II-2 above that during CSF and II-0. There were no significant differences in dry matter production and grain yield between CSF and II-0, but both were significantly greater than in the case of II-1 and II-2. Therefore, WUEi increased in the following order: II-0, CSF, II-2, II-1, although the difference was very small between II-1 and II-2. A lower crop growth rate (CGR) resulted from a decrease in the net assimilation rate (NAR) during II-1 and II-2, and there was also a reduction in the leaf area index (LAI) during II-2. Early senescence with ripening and water stress around midday decreased the rate of photosynthesis in leaves, causing the lower NAR. These physiological responses of the plants were responsible for the reduction on the dry matter production and grain yield in the intermittent irrigation.

A natural variant of NAL1 , selected in high-yield rice breeding programs, pleiotropically increases photosynthesis rate

Improvement of leaf photosynthesis is an important strategy for greater crop productivity. Here we show that the quantitative trait locus GPS (GREEN FOR PHOTOSYNTHESIS) in rice (Oryza sativa L.) controls photosynthesis rate by regulating carboxylation efficiency. Map-based cloning revealed that GPS is identical to NAL1 (NARROW LEAF1), a gene previously reported to control lateral leaf growth. The high-photosynthesis allele of GPS was found to be a partial loss-of-function allele of NAL1. This allele increased mesophyll cell number between vascular bundles, which led to thickened leaves, and it pleiotropically enhanced photosynthesis rate without the detrimental side effects observed in previously identified nal1 mutants, such as dwarf plant stature. Furthermore, pedigree analysis suggested that rice breeders have repeatedly selected the high-photosynthesis allele in high-yield breeding programs. The identification and utilization of NAL1 (GPS) can enhance future high-yield breeding and provides a new strategy for increasing rice productivity.

Hydraulic conductance as well as nitrogen accumulation plays a role in the higher rate of leaf photosynthesis of the most productive variety of rice in Japan

An indica variety Takanari is known as one of the most productive rice varieties in Japan and consistently produces 20–30% heavier dry matter during ripening than Japanese commercial varieties in the field. The higher rate of photosynthesis of individual leaves during ripening has been recognized in Takanari. By using pot-grown plants under conditions of minimal mutual shading, it was confirmed that the higher rate of leaf photosynthesis is responsible for the higher dry matter production after heading in Takanari as compared with a japonica variety, Koshihikari. The rate of leaf photosynthesis and shoot dry weight became larger in Takanari after the panicle formation and heading stages, respectively, than in Koshihikari. Roots grew rapidly in the panicle formation stage until heading in Takanari compared with Koshihikari. The higher rate of leaf photosynthesis in Takanari resulted not only from the higher content of leaf nitrogen, which was caused by its elevated capacity for nitrogen accumulation, but also from higher stomatal conductance. When measured under light-saturated conditions, stomatal conductance was already decreased due to the reduction in leaf water potential in Koshihikari even under conditions of a relatively small difference in leaf–air vapour pressure difference. In contrast, the higher stomatal conductance was supported by the maintenance of higher leaf water potential through the higher hydraulic conductance in Takanari with the larger area of root surface. However, no increase in root hydraulic conductivity was expected in Takanari. The larger root surface area of Takanari might be a target trait in future rice breeding for increasing dry matter production.

The mesophyll anatomy enhancing CO2 diffusion is a key trait for improving rice photosynthesis

Increases in rates of individual leaf photosynthesis (P(n)) are critical for future increases in yields of rice plants. Although many efforts have been made to improve rice P(n) with transgenic technology, the desired increases in P(n) have not yet been achieved. Two rice lines with extremely high values of P(n) were identified among the backcrossed inbred lines derived from the indica variety Takanari, one of the most productive varieties in Japan, and the elite japonica variety Koshihikari (Koshihikari/Takanari//Takanari). The P(n) values of the two lines at an ambient CO(2) concentration of 370μmol mol(-1) as well as at a saturating concentration of CO(2) were 20-50% higher than those of the parental varieties. Compared with Takanari, these lines had neither a higher content nor a higher activity of ribulose 1,5-bisphosphate carboxylase/oxygenase when the leaf nitrogen contents were similar, but they did have high mesophyll conductance with respect to CO(2) flux due to their higher density and more highly developed lobes of mesophyll cells. These lines also had higher electron transport rates. The plant growth rates of these lines were higher than that of Takanari. The findings show that it is possible to increase P(n) significantly, both at the current atmospheric concentration of CO(2) and at the increased concentration of CO(2) expected in the future, using appropriate combinations of genetic resources that are available at present.

Varietal Differences in Photosynthetic Rates in Rice Plants, with Special Reference to the Nitrogen Content of Leaves

Abstract The photosynthetic rate in the fl ag leaf of rice at the full heading stage was examined in three japonica varieties, Koshihikari, Aikoku and Asanohikari, and the indica high-yielding variety Takanari at the same level of leaf nitrogen. At an ambient CO2 concentration of 350 µL L-1, Takanari had a higher photosynthetic rate and stomatal conductance than the japonica varieties when plants were compared at a leaf nitrogen content of approximately 1.5 g m-2. Stomatal conductance increased considerably with increases in leaf nitrogen content in the japonica varieties. As a result, at a leaf nitrogen content of approximately 2.0 g m-2, differences in terms of the photosynthetic rate among varieties were small. By contrast, there were no clear varietal differences in Rubisco content at any identical nitrogen content of leaves. We conclude that stomatal conductance is responsible for the varietal differences in photosynthetic rate examined at the same leaf nitrogen content.

Biomass Production and Lodging Resistance in ‘Leaf Star’, a New Long-Culm Rice Forage Cultivar

Abstract Biomass production and lodging resistance in the new long-culm forage cultivar ‘Leaf Star’, developed using a precise evaluation method for lodging resistance, were evaluated by comparing these properties with those of its parents and recently improved forage rice cultivars. Leaf Star had a higher biomass production of above-ground parts than its parents, and its straw yield was 13 t ha-1. The bending moment of the basal internode at breaking in Leaf Star was three times higher than that in Koshihikari, owing to a large section modulus and a high bending stress. Biomass production of above-ground parts of Leaf Star did not differ significantly from that of other forage cultivars. However, Leaf Star had the highest straw yield of all forage cultivars. Leaf Star accumulated a large amount of starch in straw. Bending moment of the basal internode was the highest among forage cultivars owing to a large section modulus. These results show that the traits related to lodging resistance such as culm thickness and culm stiffness could be introduced into long-culm cultivars by using the precise evaluation method for the traits related to lodging resistance. The results also show that Leaf Star has a large biomass and high quality, which are suitable properties for feed and biofuel production.

Performance of a High-Yielding Modern Rice Cultivar Takanari and Several Old and New Cultivars Grown with and without Chemical Fertilizer in a Submerged Paddy Field

Abstract A high nitrogen-uptake capacity and effective use of absorbed nitrogen for dry matter and grain production are required to improve the production cost and environmental pollution. We characterized grain yield, dry matter production and nitrogen accumulation in six rice cultivars: Sekitori (released in 1848) and Aikoku (1882), referred to as SA cultivars hereafter; Koshihikari (1956); Nipponbare (1963) and Asanohikari (1987), referred to as NA cultivars hereafter; and Takanari (in 1990) as a high-yielding modern cultivar. The plants were grown with and without chemical fertilizer in a submerged paddy field. When plants were supplied with manure and chemical fertilizer, Takanari consistently produced the heaviest grain and dry matter, followed by the NA cultivars, and the SA cultivars the lightest. Dry matter production before heading was greater in Takanari and the NA cultivars due to the longer duration of vegetative growth. Dry matter production after heading was greatest in Takanari, with a larger crop growth rate (CGR), and smallest in the SA cultivars with a shorter ripening time. Greater dry matter production during ripening was accompanied by the greater accumulation of nitrogen by Takanari and NA cultivars. These plants developed a larger amount of roots. The smaller light extinction coefficient of the canopy was also attributed to the higher CGR in Takanari. When plants were grown without chemical fertilizer, Takanari also produced heavier grain and dry matter, followed by the NA cultivars. The heavier grain in these cultivars resulted from the greater dry matter production before heading, which was due to the longer period of vegetative growth. The greater dry matter production and nitrogen accumulation by Takanari and NA cultivars were evident when plants were grown with chemical fertilizer. Koshihikari was characterized by a higher CGR and greater nitrogen accumulation during ripening in the absence of chemical fertilizer which should be noted in efforts to decrease rates of nitrogen application.

Identification and characterization of genomic regions on chromosomes 4 and 8 that control the rate of photosynthesis in rice leaves

DNA marker-assisted selection appears to be a promising strategy for improving rates of leaf photosynthesis in rice. The rate of leaf photosynthesis was significantly higher in a high-yielding indica variety, Habataki, than in the most popular Japanese variety, Koshihikari, at the full heading stage as a result of the higher level of leaf nitrogen at the same rate of application of nitrogen and the higher stomatal conductance even when the respective levels of leaf nitrogen were the same. The higher leaf nitrogen content of Habataki was caused by the greater accumulation of nitrogen by plants. The higher stomatal conductance of Habataki was caused by the higher hydraulic conductance. Using progeny populations and selected lines derived from a cross between Koshihikari and Habataki, it was possible to identify the genomic regions responsible for the rate of photosynthesis within a 2.1 Mb region between RM17459 and RM17552 and within a 1.2 Mb region between RM6999 and RM22529 on the long arm of chromosome 4 and on the short arm of chromosome 8, respectively. The designated region on chromosome 4 of Habataki was responsible for both the increase in the nitrogen content of leaves and hydraulic conductance in the plant by increasing the root surface area. The designated region on chromosome 8 of Habataki was responsible for the increase in hydraulic conductance by increasing the root hydraulic conductivity. The results suggest that it may be possible to improve photosynthesis in rice leaves by marker-assisted selection that focuses on these regions of chromosomes 4 and 8.

A Comparison of the Accumulation and Partitioning of Nitrogen in Plants between Two Rice Cultivars, Akenohoshi and Nipponbare, at the Ripening Stage

Abstract To clarify the factors responsible for the maintenance of a high rate of photosynthesis at the ripening stage in the high-yield rice cultivar Akenohoshi, as compared with that in a Japanese standard cultivar, Nipponbare, we investigated the nitrogen content of leaves, focusing on the accumulation and the partitioning of nitrogen in rice plants. The nitrogen content of leaves of plants that were grown in the field or in pots remained higher in Akenohoshi than in Nipponbare during the ripening stage, and there was a close correlation between the rate of photosynthesis and the nitrogen content irrespective of cultivar and treatment. The accumulation of nitrogen in the whole plant was greater in Akenohoshi than in Nipponbare before heading and during the ripening stage. The extent of partitioning of nitrogen to leaves was higher and that to ears was lower in Akenohoshi than in Nipponbare during the ripening stage. By application of additional nitrogen fertilizer to Nipponbare, the nitrogen content of leaves was increased as a result of the increased accumulation of nitrogen in the whole plant and the enhanced partitioning of nitrogen to leaves. Our results indicate that the higher nitrogen content of Akenohoshi leaves was due to the greater accumulation of nitrogen in the plant before heading and during the ripening stage and the more effective partitioning of nitrogen to leaves during the ripening stage, which resulted in the maintenance of a high rate of photosynthesis during ripening.