Xia Ll，Xihuan KE，Tao CHEN，Yadong ZHANG，Zhen ZHP

lnstitute of Food Crops，Jiangsu AcademY of Agricultural Sciences，Jiangsu High QualitY Rice R&D Center Nanjing Branch of China National Center for Rice lmprovement，Nanjing 210014，China

ComParison among Growth，Photosynthesis and Yield in Nanjing 9108 and lts Parents during the Late DeveloPment Stage

Xia Ll*，Xihuan KE，Tao CHEN，Yadong ZHANG，Zhen ZHP

lnstitute of Food Crops，Jiangsu AcademY of Agricultural Sciences，Jiangsu High QualitY Rice R&D Center Nanjing Branch of China National Center for Rice lmprovement，Nanjing 210014，China

A good qualitY，high Yield，slow mature and middle season Japonica rice varietY，Nanjing 9108 and its parents Guandong 194 and Wuxiangjing 14，were taken as the materials for the studY.The growth and photosYnthetic characteristics of the rice plants at the post-flowering stage were compared and the rice Yield components were examined at harvest.The results showed that as compared with Wuxiangjing 14，the per plant Yield of Nanjing 9108 was 7.70%higher，and much higher than that of Guandong194.The large total grain per plant and heavY grain weight of Nanjing 9108 were the important structure basis for its Yield.Further analYsis of the growth indicators showed that Nanjing 9108 had a large leaf area of upper three green leaves area and a smaller leaf base angle in the third leaf from the bottom blades，which helped get a higher photosYnthetic capacitY in groups.On the 14thdaY after flowering，Nanjing 9108 had similar value of net photosYnthetic rate（Pn）to that of Wuxianjing 14，but lower than that of Guandong 194.Viewing from the parameters obtained from light response of photosYnthesis on the modified model of rectangular hYperbola，Pn of Nanjing 9108 was more aptated to be saturated bY light intensitY and sensitive to photoinhibition.On the whole，Nanjing 9108 with high Yield mainlY resulted from its good plant tYpe，grain structure and higher photosYnthetic capacitY in groups.Further improving its photosYnthetic capacitY of single leaf in Nanjing 9108 would be an important waY for its super high Yield potential in the future.

Japonica rice（Oryza sativa L.）；Plant tYpe；PhotosYnthetic characteristics；PhotosYnthetic light response curve；Yield

l n recent Years，Wang Cailin et al.［1］have cultivated Nanjing 9108 as a result of its parents Guandong 194 and Wuxiangjing 14 based on the selection of appearance and flavor in responsetotheshortageofnew species of Japonica rice that are of good qualitY and high Yield.ln 2013，Nanjing 9108 passed the evaluation and was considered as slow maturing japonica rice，which was suitable to be planted in Jiangsu.The Yield of such species is stable and is above 9 000 kg/hm2［1-2］.Therefore，the studies on its growth，and the species features are of great significances to the cultivation of rice in East China.NowadaYs，there are few reports，onlY a few about the cultivation of Nanjing 9108［2］and high Yield cultivation technologY［3］. ln Jiangsu，where the climate is neither too hot nor too cold，light and temperature are essential factors that affect the output of rice［4-5］.The photosYnthesis is the most essential phYsiological foundation［6］.Hence，the growth and photosYnthetic characteristics of the rice plants at the post-flowering stage were compared and the rice Yield components were examined at harvest. On the whole，Nanjing 9108 with high Yield mainlY resulted from its good plant tYpe，grain structure and higher photosYnthetic capacitY ingroups.Further improving its photosYnthetic capacitY of single leaf in Nanjing 9108 would be an important waY for its super high Yield potential in the future.

Materials and Methods

Tested materials

Nanjing 9108 and its parents Guandong 194 and Wuxiangjing 14，were taken as the materials for the studY.Nanjing 9108 was cultivated for 145 to 147 daYs，which was 3 to 5 daYs ealier than＂Huaidao No.5＂［2］.

Test design

The experiment is carried out in the field of the lnstitute of Food Crops，Jiangsu AcademY of Agricultural Sciences based on the methods proposed bY Zhu Zheng et al［7］.The crop was sowed on NaY 10，and transplanted on June 12.The distance between each strain was 20.0 cm×26.7 cm.Each species covered 9.6 m2. There were 15 lines in each area and each line included 12 strains.There were three repetitions.Five lines of crops were planted in the outside areas for protection.The soil was acid，and soil samples were collected before the experiment.The Nanjing Soil Research lnstitute of China AcademY of Agricultural Sciences determined the soil texture.The pH value was 6.20，the organics content was 1.64%，the alkaline nitrogen was 107.0 mg/kg，the available phosphorus was 36.7 mg/kg and the available potassium was 95.0 mg/kg.The experimental field can conserve water and is easY to irrigate. The nitrogen fertilizer use was 525 kg/hm2，which was applied for three times.60%fertilizer was applied 7 daYs after transplantation，and 20% fertilizer was applied 15 daYs after transplantation，and 20%fertilizer was applied during the earing period.225 kg/hm2of P2O5was applied as base fertilizer.525 kg/hm2of potassium was applied for twice，50%fertilizer on the 7thdaY after transplantation，and 50% on the mid month in JulY.

Physiological indicators and methods determination

The following growth and phYsiological indicators were determined during the earring period and different daYs after flowering.Besides，we also considered the Yield structure factors during the maturing period.

Determination of leaf base angle

Leaf base angle refers to the angle between leaves and stalk［8］.The determination was during the earring period and 14 daYs after flowering.The tillering number of each strain equaled to the five strains of tYpical plants.The leaves are intact，and grow normallY. Ten repetitions were made to each species.

Determination of leaf areas The length-width coefficient method was applied to measure the green leaf areas in this experiment.Ten spots of rice were collected from the same area，and ten leaves in the first，second and third leaf were selected randomlY.Leaf areas of the ten leaves were determined，and we calculated the total leaf areas of the last three leaves.

Determination of the stalk circumference During the grouting period，we studied 20 strains randomlY，and measured the external diameter of the main stalk with the vernier caliper.We calculated the average value of large diameter and small diameter to 0.1 mm.

The light intensity and net Photosynthetic rate curve of leaves According to Li et al［10］’s and Li et al［11］’s approach，we chose Ll-6400 convertible photosYnthetic measuring sYstem bY American Ll-COR.6400-02B LED was applied to measure the instantaneous net photosYnthetic rate. Photo flux densitY gradient was set and the net photosYnthetic rate of PFD was 0，50，100，150，200，400，600，800，1 200 and 1 400 μmol/（m2·s）.We depicted the light intensitY-Pn. Based ontheapproachbYYeand Li［12］，we learned about the maximum net photosYnthetic rate CO2（μmol/（m2·s）），the light saturation point，light compensation point and dark respiration rate CO2（μmol/（m2·s））.We calculated β and γ，and their unit was（m2·s）/μmol. The biological significance of this unit was the product of each PSll natural pigment molecule that is assimilated the area of one light quantum，and the average distance of PSll natural pigment molecule［13］.We recorded the net photosYnthetic rate when the light intensitY was within 0 and 200 μmol/（m2·s），in order to obtain AQY based on linear regression［14］.

OutPut and its comPonents Based on Jin Lei et al’s method，we selected three strains from each area and measured the strain height，the tillering number，total particle number on each strain，particle qualitY，fertilitY rate and theoretical output.

Statistics analysis

SPSS19.0 software is applied to analYze the difference significance of data and the relevance of Pearson coefficient parameters based on Duncan test through single factor variation analYsis［11］.

Results and Analyses

ComParisons of Nanjing 9108 and the comPosition factors ofits Parental outPut

According to Table 1，the actual output of ten single tillers of Nanjing 9108 was 7.70%higher than that of Wuxiangjing 14.Both were significantlY higher than that of the other parental material Guandong 194.BY analYzing its growth and Yield component，the strain height，the number of ear per plant and 1 000-grain weight increasedsignificantlY in comparison with its parental materials （P＜0.01）.The tiller of a single strain was similar to its parental strain Guandong 194，which was higherthan anotherparental Wuxiangjing 14 （P＜0.01）.However，the tiller length and fertilitY rate were significantlY lower than its two parents. Through the calculation of Yield component factors，the theoretical Yield of the total number of grains per plant of Nanjing 9108 was higher than its parents，and was even 16.69%higher than Wuxiangjing 14.Therefore，the characteristics with large sink and large grain Yield are essential structural features of Nanjing 9108，but there are still rooms to make for the Yield of Nanjing 9108.

ComParisons of leaf area of Nanjing 9108 and its Parents on the days after flowering

Relevant studies suggest that the last leaf，penultimate leaf and 3 penultimate leaf are the major contributors of Yield［15］.According to Fig.1，the leaf area of Nanjing 9108 was similar to Guandong 194，but lower than Wuxiangjing 14.The penultimate leaf was significantlY higher than its parent on the 14thdaY after flowering.The leaf area of the 3 penultimate leaves was similar to Wuxiangjing 14，and both were higherthan Guandong 194. Nanjing 9108 enjoYed a large leaf area on the 14thdaY after flowering，and was 16.69%higher than Wuxiangjing 14. Therefore，through further cultivation technologies，we maintained the green leaf area of Nanjing 9108 after flowering in order to improve its photosYnthetic rate.

ComParisons of the leaf base angle of Nanjing 9108 and its Parents at different stages

The canopY structure of directlY affects the interception of solar energY，which affects the photosYnthesis and ultimatelY influences the Yield［16］.Although the green organizations above the ground can carrY out the photosYnthesis effect，the leaf morphologY was especiallY suitable for photosYnthesis，and also lasted for the longest period［17］.The leaf base angle and canopY were especiallY close，and the leaf base angle was the main factors that determine the leaf area index［18］. According to Fig.2，compared with grouting period，the leaf base angle on the 14thdaY after flowering also increased，but there were no changes in different species.Nanjing 9108 was similar to its parental last leaf，but the penultimate leaf was distinctlY higher than Guandong 194 and similar to Wuxiangjing 14.lt is worth noticing that the leaf base angle of Nanjing 9108 was smaller than its parents.The reflection rate of 3 penultimate leaves during the tiller period was low，and its transparencY rate was high，which contributed to the interception of Nanjing 9108.

Table 2 Parameters of flag leaf photosYnthetic light response curve of Nanjing 9108 and its parents in 14thafter flowering

Changes of grains Perhilland Perimeter Per tiller in japonica Nanjing 9108 and its Parents at different stages

According to Fig.3a，Nanjing 9108 enjoYed a larger hill than its parents，and the number of grains per hill wasthe maximum on the 35thdaY after flowering.Large sink was the structural basis of its high Yield.The transportation of nutrients and moisture in the plants were largelY carried out the tube，so the number of the tube was closelY related to the transportation of nutrients and moisture.Neanwhile，the transportation of organics is an important factor to determine the Yield and qualitY of plants［19］.The stalk diameter can directlY reflect the transportation of plants.According to Fig.3b，the stalk diameter of Nanjing 9108 was dramaticallY higher than its parents Guandong 194，but was smaller than its parent Wuxiangjing 14，and there were rooms to improve to its transportation capacitY.

The resPonse curve of light intensity and Photosynthesis of jaPonica Nanjing 9108 and its Parents on the 14thday after flowering

PhotosYnthesis is the essential material foundation of crops Yield，especiallY the leaf area after flowering to determine the photosYnthesis and lightintensitY curve on the 14thdaY after flowering.According to Fig.4a，the Pn was the maximum when its parent Guandong 194 was above the middle level（＞400 μmol/（m2·s））.The Pn of Nanjing 9108 when the light intensitY was larger than 1 000 μmol/（m2·s）was higher than Wuxiangjing 14，which suggested that there was rooms for the photosYnthesis of Nanjing 9108 to improve.The changes of the evaporation rate was similar to the net photosYnthetic rate（Fig.4b）.The carboxYlation efficiencY was similar to the net photosYnthetic rate refers to the 1，5-Rubisco 1，5-RuBP in the CO2fixation process，which can be expressed bY the ratio of Pn and CO2［16，20］.According to Fig.4c，the difference of the net photosYnthetic rate of three materials was infect based on the differences in RuBP.The carboxYlation efficiencY of Guandong 194 was significantlY higher than Nanjing 9108 and Wuxiangjing 14.The carboxYlation efficiencY of Nanjing 9108 enhanced，and the value of which was between its parents.With regard to the water use efficiencY（Fig. 4d），Wuxiangjing 14 was significantlY higher than Nanjing 9108 and Guandong 194.

Based on the approaches proposed bY Ye and Li［12］，we worked out the parameters of flag leaf photosYnthetic light response curve of Nanjing 9108 and its parents in 14th after flowering（Table 2）.The apparent quantum Yield of Nanjing 9108 was lower than its parents，and the maximum net photosYnthetic rate and light saturation point were lower than its parents，which suggested that its use of high and low light intensitY did not showed anY improvement than its parents.The light compensation point and dark respiration rate were between its parents，and was significantlY lower than one of its parent WuYujing 14.γ and β of Nanjing 9108 was the largest.When γ was certain，the larger the β，the larger the light response curves.When β was certain，the larger the γ，the larger the light response curves［12-13］.Therefore，it was easY for Nanjing 9108 to saturate，and easY to have light inhibition，which was consistent with the result of low light saturation and net photosYnthetic rate.Therefore，to enhance Nanjing 9108’s response to different lights contributes to improving Yield.

Discussions

The size of canopY leaf is one of the dominant factors that affect the japonica strain height.The Japanese scholars proposed that the upper three leaves are short，thick and straight，and short leaves have high photosYnthetic efficiencY［21］.Later，Chinese scholars proposed that the photosYnthetic efficiencY of short leaves improved not because of the shortening of leaves，but because of the changes in the sources［22］.The appropriate lengthening of the upper three leaves can further enhance the japonica Yield.This paper also prove that the upper three leaves of japonica Nanjing 9108 was significantlY higher than that of Guandong 194，and the improvement of sources is the structural foundation of the high Yield.While improving the leaf area，it is necessarY to consider the spatial distribution of communitY leaves.The configuration of functional leaves and its position are essential to the ideal light state and the improvement of light use efficiencY［23］. With the addition of leaf base angle of the upper three leaves，the leaf base angle of the 3 penultimate leaves of Nanjing 9108 was dramaticallY smaller than its parents whether during the earring or tiller stage.Compared with two parents，Nanjing 9108 enjoYed the largest bank，which was the premise of high Yield.Therefore，based on the leaf position，Nanjing 9108 enjoYed a large leaf area and bank，which was an essential structure of high Yield.

Japonica fertilitY is one of the essential component factors of japonica Yield，which directlY affects the Yield potential of japonica.Studies suggested that the fertilitY rate of Nanjing 9108 was between parents，and it was lower than Wuxiangjing 14，and the theoretical Yield was higher than the actual Yield［17］.Although studies have suggested that the number of tubes of Guandong 194 and the number of airspace was less than japonica species［19］.According to the perimeter of single strain，the value of Nanjing 9108 wasdramaticallYlargerthan Guandong 194，but smaller than Wuxiangjing 14.Therefore，to ameliorate the flux contributes to the improvement of fertilitY and Yield.

Light is dYnamical power to drive photosYnthesis.Under the appropriate environmental condition，each subprocess of photosYnthesis maintained coordinated operation，which is the phYsiological experession of photosYnthetic efficiencY［4-5，24］.Based on the determination ofphotosYnthesis of Nanjing 9108 and photosYnthesis parameters，the photosYnthesis of Nanjing 9108 was smaller than Guandong 194 and similar to Wuxiangjing 14. Therefore，the large photosYnthetic advantage was not shown during the maturing period.The reduction of late stage leaves and the improvement of fertilitY rate are essential to tap the Yield potential.

ln the process of producing high-Yield and qualitY japonica，and under the premise of food qualitY，and based on the strain tYpes，the appropriate attention on the identification and selection of photosYnthetic capacitY contributes to maintaining high Yield and stable Yield.

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ResPonsible editor：Nana FAN

ResPonsible Proofreader：Xiaoyan WP

南粳9108及親本生育后期生長(zhǎng)、光合和產(chǎn)量比較

李 霞*，柯希歡，陳 濤，張亞?wèn)|，朱 鎮(zhèn) （江蘇省農(nóng)業(yè)科學(xué)院糧食作物研究所/江蘇省優(yōu)質(zhì)水稻工程技術(shù)研究中心，江蘇南京210014）

以南粳 9108及其親本為材料，研究在大田栽培條件下優(yōu)質(zhì)高產(chǎn)遲熟中粳品種南粳9108在產(chǎn)量形成的關(guān) 鍵時(shí)期的生長(zhǎng)和光合生理指標(biāo)、光強(qiáng)-光合響應(yīng)曲線，并在成熟期考察其產(chǎn)量構(gòu)成因子。結(jié)果表明：南粳9108的單株實(shí)際產(chǎn)量比親本均顯著提高，比親本武香粳14提高7.7%，大庫(kù)和大粒是其高產(chǎn)的重要結(jié)構(gòu)基礎(chǔ)。進(jìn)一步分析生長(zhǎng)指標(biāo)，南粳9108在開(kāi)花后14 d內(nèi)具有較大的綠葉面積以及倒三葉的較小葉基角，其群體的光合能力較強(qiáng)；供試材料劍葉開(kāi)花后14 d光合作用對(duì)光強(qiáng)的光合參數(shù)分析，南粳9108的光合能力與親本武香粳14類似，但顯著低于親本關(guān)東 194；而通過(guò)光合作用對(duì)光響應(yīng)的直角雙曲線修正模型分析，其對(duì)光強(qiáng)更容易飽和，且容易發(fā)生光抑制?？梢?jiàn)，高產(chǎn)水稻南粳9108具有良好的株型、穗粒結(jié)構(gòu)以及較高的群體光合能力。進(jìn)一步提高其單葉光合能力，將是其發(fā)揮超高產(chǎn)潛力的重要途徑。

粳稻；株型；光合生理特性；光強(qiáng)-光合作用曲線

國(guó)家自然科學(xué)基金項(xiàng)目（31371554）、江蘇省自主創(chuàng)新基金（CX［（12）1003］）、國(guó)家轉(zhuǎn)基因重大專項(xiàng)（2014ZX08001-004-001-009）；江蘇省自然科學(xué)基金項(xiàng)目（BK20130708）。

李霞（1970-），女，廣西桂平人，研究員，博士，主要從事植物生理育種研究，E-mail：jspplx@jaas．a(chǎn)c．cn。*通訊作者。

2015-07-09

Supported bY National Natural Scientific Fund Program （31371554）； Jiangsu lnnovation Fund （CX ［（12）1003］）； Najor National Program of Genetic Nodified Food （2014ZX-08001-004-001-009）； Jiangsu Natural Scientific Fund （BK20130708）.

*Corresponding author. E-mail: jspplx@jaas.ac.cn

Received: JulY 9， 2015 Accepted: August 10， 2015

修回日期 2015-08-10