Plant Production Science
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Print ISSN : 1343-943X
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Volume 11, Issue 1
Displaying 1-21 of 21 articles from this issue
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Review
  • Improving Drought-Avoidance Root Traits in Chickpea
    (Cicer arietinum L.) —Current Status of Research at ICRISAT
    Pooran M. Gaur, Lakshmanan Krishnamurthy, Junichi Kashiwagi
    2008 Volume 11 Issue 1 Pages 3-11
    Published: 2008
    Released on J-STAGE: January 31, 2008
    DOIhttps://doi.org/10.1626/pps.11.3
    JOURNAL FREE ACCESS
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    Chickpea (Cicer arietinum L.), an important food legume grown in the semi-arid tropical and Mediterranean regions, suffers substantial yield loss due to drought at the end of the growing season (terminal drought), as the crop is largely grown rainfed in post-rainy season on progressively receding soil moisture conditions. Root traits have been identified to postpone dehydration (drought avoidance hereafter) under moisture stress. The root length density (RLD) in the relatively shallow soil layers and the maximum root depth (RDp) were found to positively influence the seed yield under terminal drought environments. Considerable progress has been made to improve the methodology for sampling and analysis of roots. Using a PVC cylinder technique, the mini-core collection (n=211) of chickpea germplasm was evaluated for a number of root traits, including root biomass, RLD and RDp. A few germplasm accessions were identified to have a more prolific root system than the previously identified germplasm line ICC 4958, the best-known source of high root biomass. The germplasm accession ICC 8261 was identified to have the best combination of both RLD and RDp. Molecular markers have been identified for one major quantitative trait locus (QTL) that accounts for about one-third of the variation in root biomass (as measured by total root dry matter) and RDp from study of recombinant inbred lines (RILs) derived from a cross between ICC 4958 and Annigeri. New RIL populations, developed from two other crosses (ICC 8261×ICC 283 and ICC 4958×ICC 1882) involving parents having larger variation for root traits than between Annigeri and ICC 4958, are being studied to identify additional QTLs for root traits. Marker-assisted breeding for improvement of root traits in chickpea is expected to promote the development of varieties with greater drought avoidance.
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  • Genetic Opportunities to Improve Cereal Root Systems for Dryland Agriculture
    Richard A. Richards
    2008 Volume 11 Issue 1 Pages 12-16
    Published: 2008
    Released on J-STAGE: January 31, 2008
    DOIhttps://doi.org/10.1626/pps.11.12
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    Understanding the major limitations to root growth is very important if we are to maximize water and nutrient use and increase yields. Limitations may be insufficient rooting depth, root diseases, nutrient deficiencies, toxicities and soil hardness. An understanding of these limitations will lead to more precisely identifying traits for which to select and breed. Examples of successfully overcoming limiting factors to improve crop performance by breeding and selection are given for cereal cyst nematodes in wheat, soil acidity and salinity. The importance of altered crop management practices to reduce limitations is also stressed. These have resulted in a more effective and healthier root system, which results in more water use and greater yields. Opportunities to genetically increase the size of the root system in dryland systems where water and nutrients are not all used by the crop are given.
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  • Structure and Function of the Root Cap
    Morio Iijima, Shigenori Morita, Peter W. Barlow
    2008 Volume 11 Issue 1 Pages 17-27
    Published: 2008
    Released on J-STAGE: January 31, 2008
    DOIhttps://doi.org/10.1626/pps.11.17
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    The root cap (RC) is a multilayered dome of spindle-shaped parenchyma cells that overlies the growing root tip. It is present in the roots of almost all crop species. This paper briefly reviews some topics on the structure and function of the RC in the major crop species such as maize and rice. Special attention is placed on its contribution to the root system formation, that is, the elongation and growth direction of axile roots. The cells produced in the RC meristem are pushed forward as new cells form beneath them, and eventually the cells on the periphery of the RC fall off. The life cycle of RC cells of maize has been studied extensively and ranges from one to seven days. Approximately 4,000 to 21,000 cells are present in a complete maize RC, and 1,400 to 3,200 sloughed cells can be found in the rhizosphere soil per day per root. These cells, called root border cells (RBCs), mix with RC mucilage and play important roles for the root growth in soil. The RBC-mucilage complex effectively reduces the resistance roots experience during penetration into field soil, about 30-40% of the resistance being reduced by the presence of RC alone. The RC is also a tissue integral to gravitropism, and is known to determine the direction of root growth. The size of amyloplasts and coumellae in RCs has a strong influence on determining the growth angle of axile roots. The function of the individual regions of the RC and how the RC tissues and cells are formed should be studied further to advance our understanding regarding the critical roles of the RC in crop root growth.
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Crop Physiology & Ecology
  • Stomatal Responses in Rainfed Lowland Rice to Partial Soil Drying ; Evidence for Root Signals
    Joel DLC. Siopongco, Kazumi Sekiya, Akira Yamauchi, James Egdane, Abde ...
    2008 Volume 11 Issue 1 Pages 28-41
    Published: 2008
    Released on J-STAGE: January 31, 2008
    DOIhttps://doi.org/10.1626/pps.11.28
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    The role of root signals in water deficit responses of rice (Oryza sativa L.) is important in the alternate flooding and drying conditions encountered in the rainfed lowlands, where the abundant roots in shallow soil layers may generate signals when droughted, with consequent reduction in stomatal conductance (gs) and growth, despite the likelihood of additional water in deeper soil layers. This study was conducted to confirm the presence of root signals, explore their nature and plant responses, consider the suitability of the methods, and discuss implications for adaptation under rainfed lowland drought. A split-root technique was used in greenhouse studies, whereby roots were divided into two sections: flooded and droughted. The decrease in gs and transpiration rate (Tr) due to drying of a portion of the roots, and their apparent recovery upon severing of this root portion, were consistent with the role for signals. The field study confirmed the evidence for root signals during progressive soil drying, whereby gs and Tr decreased before leaf water potential (ΨL) started to decline. The increase in leaf ABA concentration under field drought, and its strong association with soil moisture tension and gs, suggested its involvement in mediating stomatal responses during early drought in rice. The recovery in ΨL after severing of droughted roots in the greenhouse could be attributed to increased hydraulic conductance. These responses imply a role for both chemical and hydraulic signals in rice, which have important implications for adaptation and crop performance in contrasting rice ecosystems.
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  • Habitat Niche-Fitness and Radix Yield Prediction Models for Angelica sinensis Cultivated in the Alpine Area of the Southeastern Region of Gansu Province, China
    Huilong Lin, Qiming Zhuang, Hua Fu
    2008 Volume 11 Issue 1 Pages 42-58
    Published: 2008
    Released on J-STAGE: January 31, 2008
    DOIhttps://doi.org/10.1626/pps.11.42
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    Dried root of Angelica sinensis has been used for thousands of years in traditional Chinese medicinal prescriptions. Researches on better knowledge of appropriate habitats for cultivation of this species are required to encourage the potential ecological sustainable industry. From 2001 to 2004, transplanting trials on the regulation of fertilizers and planting density were conducted for collection of habitat factor data at four sites of four counties in the southeastern region of Gansu Province, China. Introducing the niche theory into the research, habitat niche-fitness (HNF) is defined as the degree of similarity of an actual habitat state to the optimum habitat. A new model of HNF is constructed to evaluate the adaptive extent of A. sinensis. The results showed that the model of HNF notably outperforms the proportional similarity index and the geometric parallelism formula both in mathematical justification and biological principle testing. With HNF as a surrogate for composite environmental factors, a radix yield model was constructed. Evaluation of the present model by the sampled subplots specified for data validation proved that the model could be well used for predicted of radix yield across a wide-spread area. The radix yield prediction model and its uses are recommended within the limitations of the data used in the study area. Beyond this range, validation of the radix yield prediction model will be necessary.
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  • Alternative Respiratory Pathway under Drought is Partially Mediated by Hydrogen Peroxide and Contributes to Antioxidant Protection in Wheat Leaves
    Hanqing Feng, Jiangong Duan, Hongyu Li, Houguo Liang, Xin Li, Na Han
    2008 Volume 11 Issue 1 Pages 59-66
    Published: 2008
    Released on J-STAGE: January 31, 2008
    DOIhttps://doi.org/10.1626/pps.11.59
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    Water stress significantly enhanced the capacity of alternative respiratory pathway and induced AOX1 transcript in wheat (Triticum aestivum L.) leaves. The water-stressed seedlings pretreated with 1 mM salicylhydroxamic acid (SHAM) had higher level of production of reactive oxygen species (ROS) than the seedlings either subjected to drought or SHAM treatment alone did. This observation suggests that cyanide-resistant respiration could play a role in antioxidant protection under the condition of drought. Exogenous application of hydrogen peroxide effectively increased the capacity of alternative respiratory pathway and induced AOX1 transcription. Pretreatment with ROS scavengers, such as 4,5-dihydroxy-1,3-benzene disulfonic acid (Tiron) and dimethylthiourea (DMTU), arrested the increase of ROS and partly inhibited the induction of both cyanide-resistant respiration and AOX1 transcript under water stress. These results suggest that the enhancement of cyanide-resistant respiration under drought might be partially mediated by hydrogen peroxide.
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  • Rice Cultivars with Higher Sucrose Synthase Activity Develop Longer Coleoptiles under Submerged Conditions
    Akari Fukuda, Satoshi Yoshinaga, Kenji Nagata, Hiroyuki Shiratsuchi
    2008 Volume 11 Issue 1 Pages 67-75
    Published: 2008
    Released on J-STAGE: January 31, 2008
    DOIhttps://doi.org/10.1626/pps.11.67
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    Sucrose synthase, which catalyzes uridine diphosphate (UDP)-dependent cleavage of sucrose into fructose and UDP-glucose, is induced by oxygen deficiency in rice seedlings and is considered to play an important role in energy production under hypoxic conditions. In this study, we analyzed the relationship between coleoptile elongation and sucrose synthase activity in rice (Oryza sativa L.) cultivars under submerged conditions. We also analyzed the activity of α-amylase, which digests starch reserves in the endosperm and is considered to be important for energy production in young seedlings. The results indicated that different rice cultivars had different sucrose synthase and α-amylase activities under submerged conditions. Moreover, sucrose synthase activity in whole seedlings was significantly correlated with coleoptile length under submerged conditions, whereas the correlation between α-amylase activity and coleoptile length was low. Sugar content of shoots differed with the cultivar. Correlation analysis demonstrated that sucrose content was highly correlated with coleoptile length and sucrose synthase activity, but not with α-amylase activity.
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  • Developmental Responses of Wheat cv. Norin 61 to Fluence Rate of Green Light
    Shin-ya Kasajima, Naoto Inoue, Rezwanul Mahmud, Masakazu Kato
    2008 Volume 11 Issue 1 Pages 76-81
    Published: 2008
    Released on J-STAGE: January 31, 2008
    DOIhttps://doi.org/10.1626/pps.11.76
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    Wheat (Triticum aestivum L. cv. Norin 61) plants were grown under five different photon flux densities obtained by modulating the number of green light-emitting diodes (LEDs) under white fluorescent lamps. The experiment was conducted under continuous irradiation at a constant temperature of 20°C to clarify the developmental responses of wheat to the fluence rate of green light. The higher the photon flux density of green light, the shorter the number of days from emergence to heading. The earliest heading was observed in the plants grown under 496 green LEDs, 32.0 days after emergence. A significant logarithmic function could fit the relationship between the fluence rate of green light and developmental rate. In this report, principal component analysis (PCA) was adopted to analyze the confounding of green light versus photosynthetic photon flux density (PPFD) with 17 developmental and morphological traits. The eigenvalues explained were 5.64, 3.20, and 2.61, respectively, for the first, second and third principal components (PCs). The first PC was assumed as the factor related to the isometric growth, and the third PC was assumed as the factor related to the developmental rate and culm elongation. Therefore, it was supposed that the first and third PCs were affected by the PPFD and the photon flux density of green light, respectively. The results suggested that the fluence rate of green light affects the development of wheat as a signal source. Furthermore, the development of wheat was promoted by the green light independent of PPFD.
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  • Evaluation of Cultivar Differences in Preharvest Sprouting of Common Buckwheat (Fagopyrum esculentum Moench)
    Takahiro Hara, Takahisa Tetsuka, Katsuhiro Matsui, Hiroki Ikoma, Akira ...
    2008 Volume 11 Issue 1 Pages 82-87
    Published: 2008
    Released on J-STAGE: January 31, 2008
    DOIhttps://doi.org/10.1626/pps.11.82
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    Preharvest sprouting of buckwheat (Fagopyrum esculentum Moench) is an important problem, but cultivar differences in preharvest sprouting and their causes have not been investigated. We detected cultivar differences under natural field conditions. Preharvest sprouting of three cultivars was significantly lower than that of the current main cultivar. Seeds collected before rainfall were threshed and incubated on a wet filter paper in a petri dish for 10 days at 10, 20, 30 and 40°C in the dark, or at an alternating light and temperature condition of 8 h light at 30°C and 16 h darkness at 20°C. Germination was promoted by a higher temperature except for 40°C, suggesting that the risk of preharvest sprouting in buckwheat is higher at a relatively higher temperature. The risk of preharvest sprouting in the field was highly correlated with germination at 20°C (r = 0.98***) and 30°C (r = 0.99***) in the dark, suggesting that germination test can be used to predict preharvest sprouting in the field. Preharvest sprouting was significantly correlated (r = -0.77**) with main stem length, suggesting that ecotype is partly responsible for this phenomenon.
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  • Transient Expression of Green Fluorescent Protein in Rice Calluses : Optimization of Parameters for Helios Gene Gun Device
    Nono Carsono, Tomohiko Yoshida
    2008 Volume 11 Issue 1 Pages 88-95
    Published: 2008
    Released on J-STAGE: January 31, 2008
    DOIhttps://doi.org/10.1626/pps.11.88
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    An optimized condition for particle bombardment is necessary for efficient genetic transformation. Parameters for Helios gene gun, the new system for nucleic acid delivery which is mainly consists of hand-held device sold by Bio-Rad Laboratories (California USA), were examined based on transient expression of synthetic green fluorescent protein (sgfp) in rice calluses of indica cv. Fatmawati and japonica cv. Nipponbare. In the experimental conditions that we examined, parameters found to be the most favorable conditions for transient expression of sgfp in rice callus cells were as follows: 200-250 psi helium pressure, 0.6 μm gold particle size, 0.25 mg gold particles per shot, and 1.5 μg plasmid-DNA per shot. Desiccation of callus cells for eight min was also found appropriate. The level of transient sgfp expression was not significantly influenced by the pre-culture for 4 to 12 d before bombardment or by callus age between 10 and 33 wk old in Fatmawati. These parameters for this particular device should improve the transient expression, thus enabling stable expression of introduced genes via Helios gene gun using callus as a target tissue.
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  • Yield Stability of Aromatic Upland Rice with High Yielding Ability in Indonesia
    Totok Agung Dwi Haryanto, Suwarto, Tomohiko Yoshida
    2008 Volume 11 Issue 1 Pages 96-103
    Published: 2008
    Released on J-STAGE: January 31, 2008
    DOIhttps://doi.org/10.1626/pps.11.96
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    Aromatic rice variety, Mentikwangi, was crossed with high-yielding upland rice variety, Poso, and the pedigree was selected to obtain lines with high yielding and aromatic characters. The objectives of the research were to study the yield stability of aromatic upland genotypes across different locations and to select aromatic upland rice genotypes having wide adaptability, and or specific location adaptability. Yield stability of genotypes was estimated by using regression lines proposed by Finlay and Wilkinson. Some genotypes showed high yield stability and wide adaptability in different locations, and others showed good adaptability to a specific location. The lines having high yield stability and wide adaptability were G10 (405 g m-2), G19 (400 g m-2), G39 (418 g m-2), and G136 (411 g m-2), which may be considered as candidates of new aromatic upland rice cultivars. Situpatenggang had specific adaptability at the fertile locations; and Poso and G13 at the infertile locations. Genotype x location interactions for the yield and its components performance were observed.
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  • Overexpression of a Maize SPS Gene Improves Yield Characters of Potato under Field Conditions
    Ken Ishimaru, Naoki Hirotsu, Takayuki Kashiwagi, Yuka Madoka, Kiyoshi ...
    2008 Volume 11 Issue 1 Pages 104-107
    Published: 2008
    Released on J-STAGE: January 31, 2008
    DOIhttps://doi.org/10.1626/pps.11.104
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    We analyzed the yield characters of field-grown transgenic potato plants (Solanum tuberosum) carrying a maize gene for sucrose-phosphate synthase (SPS), the key enzyme in sucrose synthesis. The SPS activity in the leaves of transgenic plants (line Ag1203) was 2 times that of the control (cv. May Queen). There was no difference in the photosynthetic CO2 uptake rates between Ag1203 and May Queen plants, and the leaf starch content of Ag1203 was lower. These observations indicate that the introduction of a foreign SPS gene improved the supply of photosynthate from source (leaves) to sink (tubers). Additionally, leaf senescence of the transgenic potato plants was delayed relative to that of May Queen. The average tuber weight and total yield of Ag1203 plants were at least 20% higher, and the tuber sucrose content, which is related to eating quality, was also higher. Increased translocation of photosynthate and longer period of photosynthetic activity in the leaves may have increased the yield of Ag1203. These results suggest that introduction of the SPS gene improved the yield characters and quality of potato tubers under field conditions.
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  • Development of Rice “Seed-Mats” Consisting of Hardened Seeds with a Cover of Soil for the Rice Transplanter
    Hiroyuki Shiratsuchi, Hisashi Kitagawa, Kensuke Okada, Kazuyasu Nakani ...
    2008 Volume 11 Issue 1 Pages 108-115
    Published: 2008
    Released on J-STAGE: January 31, 2008
    DOIhttps://doi.org/10.1626/pps.11.108
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    Conventional seedling mats for rice transplanters in Japan are heavy to carry, and much labor is required to collect, wash and store the nursery boxes. In addition, seeding time overlaps with the labor peak in spring. To reduce such labor, we developed a “seed-mat” consisting of hardened rice seeds (Oryza sativa L.) glued onto a molded rice-hull mat with a cover of soil glued on. Seed hardening, which is done by soaking seeds in water at 15°C for 5 d followed by drying, and heating of the seeds at 50°C for 5 or 7 d, reduced the time to 50% germination (T50). In dormant seeds, the heating before the hardening enhanced the effect of the hardening more than the heating after the hardening. Seed hardening increased the dry weight of the shoots from the seed-mats. Covering the mat with soil also increased the percentage of seedling emergence, shoot length, number of leaves and shoot dry weight of the seed-mats. The hardened seeds maintained short T50 and 95% or higher germination for 120 d at room temperature. In the seed-mat, the hardened seeds maintained 95% or higher seedling emergence for 208 d. In conclusion, superior seed-mats could be produced using the procedure involving both seed hardening and cover of soil. The seed-mats can be prepared in winter and stored until seedling-raising period in spring. By the seed-mats, seedling mats can be prepared for the rice transplanter without using nursery boxes, which take up storage space and require much labor.
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  • Role of Belowground Parts of Green Manure Legumes, Crotalaria spectabilis and Sesbania rostrata, in N Uptake by the Succeeding Tendergreen Mustard Plant
    Bongsu Choi, Masamichi Ohe, Jiro Harada, Hiroyuki Daimon
    2008 Volume 11 Issue 1 Pages 116-123
    Published: 2008
    Released on J-STAGE: January 31, 2008
    DOIhttps://doi.org/10.1626/pps.11.116
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    Using symbiotically N2-fixing legumes as green manures is a way to supply N from the atmosphere to cropping ecosystems. Usually whole plants of the green manure are incorporated into soil; hence, the belowground parts as well as the aboveground parts would contribute to N transfer to succeeding crops. However, little is known about the contribution of the belowground parts alone. We assessed N transfer from belowground parts compared to whole plants of two legumes, Crotalaria spectabilis and Sesbania rostrata. Each of the legumes was grown approximately for 3 months in a 1/2000a Wagner pot filled with soil media, and then the roots alone (R) or shoot and root (S+R) were harvested and incorporated in the pots. Tendergreen mustard (Brassica rapa) as the succeeding crop was grown for 66 days in these pots without additional fertilizer. Although the amount of N in green manure in S+R pots was approximately 4-fold higher than that in R pots, differences in N uptake by tendergreen mustard between the S+R and R pots were smaller (1.7-fold for C. spectabilis and 2.3-fold for S. rostrata). This means that N recovery rate by tendergreen mustard was significantly higher in R than in S+R pots with either green manures. Differences in C/N ratio of the green manures could not likely explain the higher N recovery rate in R pots. Bioassay of the aqueous extracts from the green manure with lettuce seedlings suggested that growth inhibitory effects might be responsible for the lower recovery rate in S+R treatment.
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(Short Report)
Genetic Resources Evaluation
  • Responses of Rice Genotypes Carrying Different Dwarf Genes to Fusarium moniliforme and Gibberellic Acid
    Liangyong Ma, Zhijuan Ji, Jinsong Bao, Xudong Zhu, Ximing Li, Jieyun Z ...
    2008 Volume 11 Issue 1 Pages 134-138
    Published: 2008
    Released on J-STAGE: January 31, 2008
    DOIhttps://doi.org/10.1626/pps.11.134
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    A total of 32 rice genotypes carrying different dwarf or semi-dwarf genes were inoculated with the fungus Fusarium moniliforme Sheldon or treated with 50 mg l-1 GA3 in order to select resources resistant to rice bakanae disease from the dwarf materials. The length of the elongated seedlings was measured, and the percentage of death of the seedlings after transplanting to field was also counted. A significant correlation was found between the length of the seedling treated with GA3 and disease injury by bakanae fungus. Rice materials carrying dwarf gene such as sd1 were not only sensitivity to GA3 but also susceptive to rice bakanae disease. Materials carrying dwarf gene d1 were insensitive to GA3 but susceptive to bakanae. On the other hand, all materials carrying d29, sd6 or sdq(t) genes showed resistance to bakanae. The present study indicated that dwarf and semi-dwarf rice materials might be useful resources for improvement of bakanae resistance in rice breeding programs.
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Crop Morphology
  • Correlation between Chloroplast Ultrastructure and Chlorophyll Fluorescence Characteristics in the Leaves of Rice (Oryza sativa L.) Grown under Salinity
    Koji Yamane, Michio Kawasaki, Mitsutaka Taniguchi, Hiroshi Miyake
    2008 Volume 11 Issue 1 Pages 139-145
    Published: 2008
    Released on J-STAGE: January 31, 2008
    DOIhttps://doi.org/10.1626/pps.11.139
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    The seedlings of Oryza sativa L. cv. Nipponbare grown by hydroponic culture for 3 wks were treated with 75, 100, 150 and 200 mM NaCl for 14, 14, 6 and 3 days, respectively, and examined for chloroplast ultrastructure in the region where chlorophyll fluorescence had been recorded. NaCl treatment decreased the ratio of variable to maximum chlorophyll fluorescence yield (Fv/Fm) and caused swelling of thylakoids. The swelling of thylakoids was quantified by the percentage of the length of swollen thylakoids to the total length of thylakoids. This value was increased with increasing NaCl concentration. Although Fv/Fm decreased at all concentrations of NaCl, the minimal fluorescence yield F0 was not increased by the treatment with 75 or 100 mM NaCl. The percentage of the length of swelling was low at 75 and 100 mM NaCl. On the other hand, F0 increased and the swelling of thylakoids was prominent with 150 and 200 mM NaCl treatment. These results suggest that the decrease in Fv/Fm due to the increase in F0 under salt stress correlates with the ultrastructural damage. The decrease in Fv/Fm due to the increase in F0 is expected to be useful as an indicator to evaluate the damage in chloroplasts, especially in thylakoid membranes, under salinity.
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Post Harvest Physiology
(Short Report)
Agronomy
  • Effect of Nitrogen Application Rate and Timing on Grain Yield and Protein Content of the Bread Wheat Cultivar ‘Minaminokaori’ in Southwestern Japan
    Hiroshi Nakano, Satoshi Morita, Osamu Kusuda
    2008 Volume 11 Issue 1 Pages 151-157
    Published: 2008
    Released on J-STAGE: January 31, 2008
    DOIhttps://doi.org/10.1626/pps.11.151
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    The effect of nitrogen (N) application rates at active tillering and anthesis on grain yield and protein content of a bread wheat cultivar, ‘Minaminokaori’, was examined in the field study conducted in Fukuoka, Japan. Number of spikes, leaf area index (LAI), and SPAD value at anthesis increased significantly (P<0.05) with increasing N application rate at active tillering. Grain yield also increased significantly (P<0.05) because of the increased number of spikes. However, grain yield did not increase significantly (P>0.05) with increasing N application rate at anthesis. The slope of the relationship between N application rate at active tillering and grain yield was about 3 times that of the relationship between N application rate at anthesis and grain yield. These results indicate that N application at active tillering is more effective than N application at anthesis for increasing grain yield. Increasing N application rate at active tillering and anthesis generally increased grain protein content. However, the slope of the relationship between N application rate at anthesis and grain protein content was about 2 times that of the relationship between N application rate at active tillering and grain protein content. These results indicate that N application at anthesis is more effective than N application at active tillering for increasing grain protein content. The interaction between N application rates at active tillering and anthesis was significant (P<0.05) for grain protein content. With the application of 4 g N m-2 at active tillering, grain protein content increased linearly at a rate of about 0.5% per 1 g N m-2 (from 10.9% to 14.0%) with increasing N application rate (from 0 to 6 g N m-2) at anthesis. However, the rate of increase in grain protein content with increasing N application rate at anthesis was greater with 0 g N m-2 at active tillering than that with 4 g N m-2 at active tillering, whereas that with 8 g N m-2 at active tillering was smaller than that with 4 g N m-2 at active tillering. Application of 8 g N m-2 at tillering resulted in the highest SPAD value at anthesis; this was followed by the results for 4 and 0 g N m-2. Therefore, the SPAD value may be an important trait to decide N application rate at anthesis.
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