嫁接番茄果实糖分变化及其与氮磷含量相关性分析

doi:10.16420/j.issn.0513-353x.2023-0850

摘要/Abstract

摘要：

以红果番茄‘戴维森’（Dv）和粉果番茄‘罗拉’（La）为接穗，以‘Anta-T’（At）和‘Midako’（Md）番茄为砧木进行嫁接，研究了不同穗砧组合不同发育时期果实糖组分含量、蔗糖代谢相关酶活性及矿质元素含量的变化。结果表明，砧木可显著影响接穗果实的果糖、葡萄糖和蔗糖含量且与接穗间存在显著交互作用。花后30 d，Dv/At果实蔗糖含量较Dv/Md高8.06%，La/At和La/Md之间无显著差异。花后110 d时，Dv/Md果糖含量较自根嫁接苗低20.66%，La/Md却与自根嫁接苗无显著差异。不同嫁接组合果实中糖分含量差异与蔗糖代谢酶活性密切相关，果实糖分差异在发育前期主要受到接穗对酸性转化酶活性的调控作用，发育后期主要受到砧木对蔗糖磷酸合成酶和中性转化酶活性的影响。不同穗砧组合的果实中氮、磷、钾、钙、镁和铁含量差异显著。曲线估计分析显示，果实中氮和磷含量与蔗糖酸性转化酶和中性转化酶活性以及果糖、葡萄糖含量呈显著的负相关，与蔗糖含量成显著正相关。因此，在嫁接栽培促进番茄提质增产过程中，氮、磷养分管理应与砧木吸收特性相协调，从而避免氮、磷吸收增多引起番茄果实糖度和风味降低。

关键词: 番茄, 砧木, 接穗, 糖分积累, 氮磷

Abstract:

The changes of sugar content，sucrose metabolism related enzyme activities and mineral elements in tomato fruits at different developmental stages were studied. Red fruit tomato‘Davis’(Dv) and pink fruit tomato‘Rolla’(La) were used as scions，and tomato‘Anta-T’(At) and‘Midako’(Md) were used as rootstocks. Regulation of the content of fructose，glucose and sucrose by scion and rootstock in tomato fruits at different developmental stages were found，and there was a significant interaction between scion and rootstock. At 30 d after flowering，sucrose content in fruit of Dv/At was 8.06% higher than that of Dv/Md，but there was no significant difference between La/At and La/Md. Meanwhile fructose content (16.44 mg · g^-1FW) in Dv/Md was 20.66% lower than Dv/Dv at 110 d after flowering，but there was no significant difference between La/Md and La/La. The difference of sugar content in tomato fruit with different grafting combinations was closely related to the activity of sucrose metabolic enzyme. The difference of sugar content in the early stage of fruit development of grafted tomato was mainly regulated the activity of acid invertase (AI) by scion in fruit. The difference in sugar content at late development stage was mainly due to the influence of rootstock on the activity of sucrose phosphate synthetase (SPS) and neutral invertase (NI). At the same time，nitrogen (N)，phosphorus (P)，potassium (K)，calcium (Ca)，magnesium (Mg) and iron (Fe) in tomato fruits with different grafting combinations were significantly different. Curve estimation analysis showed that N and P content in fruit were significantly negatively correlated with invertase activity (AI and NI)，fructose and glucose，and significantly positively correlated with sucrose content. Therefore，in the process of tomato grafting to promote the quality and yield of tomato，the management of nitrogen and phosphorus nutrients after grafting should be coordinated with the absorption characteristics of rootstocks，so as to avoid the decrease of sugar content and flavor of tomato fruits caused by increase of nitrogen and phosphorus absorption.

Key words: tomato, rootstock, scion, sugar accumulation, nitrogen and phosphorus

王磊, 高方胜, 张志焕, 李炜蔷, 曹逼力, 徐坤. 嫁接番茄果实糖分变化及其与氮磷含量相关性分析[J]. 园艺学报, 2024, 51(2): 361-371.

WANG Lei, GAO Fangsheng, ZHANG Zhihuan, LI Weiqiang, CAO Bili, XU Kun. Change of Sugar in Grafted Tomato Fruit and Correlation Analysis with Nitrogen and Phosphorus Content[J]. Acta Horticulturae Sinica, 2024, 51(2): 361-371.

图/表 9

参考文献 38

[1]	Aloni B, Cohen R, Karni L, Aktas H, Edelstein M. 2010. Hormonal signaling in rootstock-scion interactions. Scientia Horticulturae, 127:119-126.
[2]	Bharali A, Baruah K K. 2022. Effects of integrated nutrient management on sucrose phosphate synthase enzyme activity and grain quality traits in rice. Physiol Mol Biol Plants, 28 (2):383-389.
[3]	Coura F T, Fries D D, Quoos R D, Teixeira F A, Pires A J, Figueiredo A J. 2020. Activity invertase and amylase in Marandu grass under shading and nitrogen fertilization. Acta Scientiarum Agronomy, 42:e42496.
[4]	Gao Fang-sheng, Wang Lei, Xu Kun. 2014. Comprehensive evaluation of relationship between rootstocks and yield and quality in grafting tomato. Scientia Agricultura Sinica, 47 (3):605-612. (in Chinese)
	高方胜, 王磊, 徐坤. 2014. 砧木与嫁接番茄产量品质关系的综合评价. 中国农业科学, 47 (3):605-612. doi: 10.3864/j.issn.0578-1752.2014.03.020
[5]	Gong T, Brecht G K, Hutton S F, Koch K E, Zhao X. 2022. Tomato fruit quality is more strongly affected by scion type and planting season than by rootstock type. Frontiers in Plant Science, 13:948556.
[6]	He Wen, Pan He-li, Pan Teng-fei, Tang Hao-ru, Wang Xiao-rong, Pan Dong-ming. 2017. Research progress on the interaction between scion an rootstock in fruit trees. Acta Horiculturae Sinica, 44 (9):1645-1657. (in Chinese)
	何文, 潘鹤立, 潘腾飞, 汤浩茹, 王小蓉, 潘东明. 2017. 果树砧穗互作研究进展. 园艺学报, 44 (9):1645-1657. doi: 10.16420/j.issn.0513-353x.2017-0147
[7]	Leonardi C, Giuffrida F. 2006. Variation of plant growth and macronutrient uptake in grafted tomatoes and eggplants on three different rootstocks. European Journal of Horticultural Science, 71 (3):97-101.
[8]	Li Fan, Wei Hua, Qi Jian-guo, Sun Si-min, Zou ang-jun, Li Ming-jun. 2021. Effects of nitrogen application at mature stage on sugar contents and related gene expression of Fuji apple. Journal of Northwest A & F University, 49 (8):111-119. (in Chinese)
	李凡, 魏桦, 戚建国, 孙思敏, 邹养军, 李明军. 2021. 成熟期试氮对富士苹果糖含量及相关基因表达的影响. 西北农林科技大学学报(自然科学版), 49 (8):111-119.
[9]	Liu De-xing, Jing Xin, Jiao Juan, Wei Min, Sui Shen-li, Zhao Li-hua, Li Yan-wei, Zhao Na, Gong Biao, Shi Qing-hua. 2017. Comprehensive evaluation of yield,quality and salt stress tolerance in grafting tomato. Acta Horticulturae Sinica, 44 (6):1094-1104. (in Chinese)
	刘德兴, 荆鑫, 焦娟, 魏珉, 隋申利, 赵利华, 李艳玮, 赵娜, 巩彪, 史庆华. 2017. 嫁接对番茄产量、品质及耐盐性影响的综合评价. 园艺学报, 44 (6):1094-1104. doi: 10.16420/j.issn.0513-353x.2016-0947
[10]	Liu Hui-ying, Zhu Zhu-jun, Qian Qiong-qiu, Ge Zhi-ping. 2004. The effects of different rootstocks on the sugar metabolism and related enzyme activities in small and early-maturing watermelon during fruit development. Acta Horticulurae Sinica, 31 (1):47-52. (in Chinese)
	刘慧英, 朱祝军, 钱琼秋, 葛志平. 2004. 砧木对小型早熟西瓜果实糖代谢及相关酶活性的影响. 园艺学报, 31 (1):47-52.
[11]	Liu Xiang-yu, Li Juan, Huang Min, Liang Chun-hui, Chen Jie-zhong. 2015. Research on influences of rootstock on sugar accumulation in ‘Shatangju’Tangerine fruits. Scientia Agricultura Sinica, 48 (11):2217-2228. (in Chinese) doi: 10.3864/j.issn.0578-1752.2015.11.013
	刘翔宇, 李娟, 黄敏, 梁春辉, 陈杰忠. 2015. 柑橘砧木对砂糖橘果实糖积累的影响. 中国农业科学, 48 (11):2217-2228. doi: 10.3864/j.issn.0578-1752.2015.11.013
[12]	Osorio S, Ruan Y L, Fernie A R. 2014. An update on source-to-sink carbon partitioning in tomato. Frontiers in Plant Science, 5:1-11.
[13]	Paula B V, Sete P B, Berghetti L P, Silva L O, Jung J P, Nicoloso F T, Mayer N A, Kulmann M S, Brunetto G. 2022. Kinetic parameters related to nitrogen uptake in‘Okinawa’peach rootstocks are altered by‘Chimarrita’scion Nitrogen uptake in‘Okinawa’peach rootstock. Journal of the Science of Food and Agriculture, 103 (2):917-923.
[14]	Qi Hong-yan, Jiang Yan-yan, Hua Li-jing. 2012. Responses to short-term low night temperature of sucrose-metabolizing in domesticated tomato and wild relative. Acta Horticulturae Sinica, 39 (2):281-288. (in Chinese)
	齐红岩, 姜岩岩, 华利静. 2012. 短期夜间低温对栽培番茄和野生番茄果实蔗糖代谢的影响. 园艺学报, 39 (2):281-288.
[15]	Qi Hong-yan, Li Tian-lai, Zhang Jie, Liu Hai-tao. 2006. Relationship between carbohydrate change and related enzymes activities during tomato fruit development. Acta Horticulturae Sinica, 33 (2):294-299. (in Chinese)
	齐红岩, 李天来, 张洁, 刘海涛. 2006. 番茄果实发育过程中糖的变化与相关酶活性的关系. 园艺学报, 33 (2):294-299.
[16]	Qin Wei, Cui Xin-yi, Chen Bo-lang, Li Jian-gui. 2013. Effects of different fertilization ratio of N,P and K on related enzyme activities of sugar accumulation in Xinjiang Red Fuji apple. Journal of Xinjiang Agricultural University, 36 (6):431-436. (in Chinese)
	秦伟, 崔新仪, 陈波浪, 李建贵. 2013. 不同氮磷钾配比施肥对新疆红富士苹果糖分积累相关酶活性的影响. 新疆农业大学学报, 36 (6):431-436.
[17]	Ru L, He Y, Zhu Z J, Patrick J W, Ruan Y L. 2020. Integrating sugar metabolism with transport:elevation of endogenous cell wall invertase activity up-regulates SIHT2 and SISWEET12c expression for early fruit development in tomato. Frontiers in Genetics, 11:592596.
[18]	Savvas D, Oztekin G B, Tepecik M, Ropokis A, Tuzel Y. 2017. Impact of grafting and rootstock on nutrient-to-water uptake ratios during the first month after planting of hydroponically grown tomato. Journal of Horticultural Science and Biotechnology, 92 (3):1-9.
[19]	Stella K, Maina M, Jesca M. 2020. Identification of Ralstonia solanacearum resistant rootstocks for tomato grafting. Journal of Animal & Plant Sciences, 43 (3):7452-7457.
[20]	Shang Le-Le, Song Jian-wen, Wang Jia-ying, Zhang Yu-yang, Ye Zhi-biao. 2019. Research progress on quality formation and molecular mechanism of tomato fruit. China Vegetables,(4):21-28. (in Chinese)
	尚乐乐, 宋建文, 王嘉颖, 张余洋, 叶志彪. 2019. 番茄果实品质形成及其分子机理研究进展. 中国蔬菜,(4):21-28.
[21]	Sun N, Ma Y P, Wang X J, Ying Q S, Huang Y P, Zhang L C, Zhu Z J, Wang Y H, He Y. 2023. Grafting onto pumpkin alters the evolution of fruit sugar profile and increases fruit weight through invertase and sugar transporters in watermelon. Scientia Horticulturae, 314:11936.
[22]	Tang G Q, Luscher M, Sturm A. 1999. Antisense repression of vacuolar and cell wall invertase in transgenic carrot alters early plant development and sucrose partitioning. Plant Cell, 11 (2):177-189. doi: 10.1105/tpc.11.2.177 pmid: 9927637
[23]	Tecchio M A, Silva M J, Callili D, Angeli G M, Silva D A, Neto F J, Moura M F. 2019. Nutrient uptake in‘Venus’grape vine grafted onto different rootstocks during two growing seasons. Australian Journal of Crop Science, 13 (10):1594-1599.
[24]	Tian H, Ma L, Cong Z, Hao H, Gong B, Yu X, Wang X. 2010. Antisense repression of sucrose phosphate synthase in transgenic muskmelon alters plant growth and fruit development. Biochemical and Biophysical Research Commtmications, 393 (3):365-370.
[25]	Turhan A, Ozmen N, Kuscu H, Serbeci M S, Seniz V. 2012. Influence of rootstocks on yield and fruit characteristics and quality of watermelon. Horticulture Environment Biotechnology, 53 (4):336-341.
[26]	Turhan A, Ozmen N, Serbeci M S, Seniz V. 2011. Effects of grafting on different rootstocks on tomato fruit yield and quality. Horticultural Science, 38 (4):142-149.
[27]	Wang J W, Li H J, Wang Q, Huang X L, Hu W, Wang S S, Zhou Z G. 2023. Effects of phosphorus application on carbohydrate metabolism in cottonseed kernel during the key development period provided a new insight for phosphorus management in cotton production. Industrial Crops and Products, 191:115972.
[28]	Wang Lei, Gao Fang-sheng, Xu Kun. 2017. Effect of rootstock-scion interaction on plant growth and leaf carbon-nitrogen assimilation in overwintering tomato. Plant Physiology Journal, 53 (9):1695-1702. (in Chinese)
	王磊, 高方胜, 徐坤. 2017. 砧穗互作对越冬番茄生长及叶片碳氮同化能力的影响. 植物生理学报, 53 (9):1695-1702.
[29]	Wei Jie, Jiang Yuan, Lin Cai-xia, Shi Lei, Luo Tao-feng, Li Yong-feng. 2019. Correlation and path analysis between fruit quality and mineral elements of 6 varieties of Korla Fragrant Pear. Food Science, 40 (4):259-265. (in Chinese) doi: 10.7506/spkx1002-6630-20180306-070
	位杰, 蒋媛, 林彩霞, 石蕾, 罗洮峰, 李永丰. 2019. 6个库尔勒香梨品种果实矿质元素与品质的相关性和通径分析. 食品科学, 40 (4):259-265. doi: 10.7506/spkx1002-6630-20180306-070
[30]	Xu Chuan-qiang, Li Tian-lai, Qi Hong-yan. 2006. Effect of grafting on development,carbohydrate content and sucrose-metabolizing enzymes activities of muskmelon fruit. Acta Horticulturae Sinica, 33 (4):773-778. (in Chinese)
	许传强, 李天来, 齐红岩. 2006. 嫁接对网纹甜瓜果实发育、糖含量及蔗糖代谢相关酶活性的影响. 园艺学报, 33 (4):773-778.
[31]	Xu Sheng-li, Chen Qing-yun, Li Shao-hua, Zhang Li-li, Gao Jiang-sheng, Wang He-li. 2005. Role of sugar-metabolizing enzymes and GA₃,ABA in sugars accumulation in grafted muskmelon fruit. Journal of Fruit Science, 22 (5):514-518. (in Chinese)
	徐胜利, 陈青云, 李绍花, 张利莉, 高疆生, 王合理. 2005. 糖代谢相关酶和GA₃、ABA在嫁接伽师瓜果实糖分积累中的作用. 果树学报, 22 (5):514-518.
[32]	Yu Xin-jian. 1985. Experimental manual of plant physiology. Shanghai: Shanghai Science And Technology Press:148-149. (in Chinese)
	於新建. 1985. 植物生理学实验手册. 上海: 上海科学技术出版社:148-149.
[33]	Yuan Ting-ting, Song Xiao-yi, Wang Zhong-bin, Yang Jian-ping, Xu Kun. 2011. Effect of grafting cultivation and fertilization on the yield,NPK uptake and utilization of tomatoes. Plant Nutrition and Fertilizer Science, 17 (1):131-136. (in Chinese)
	袁亭亭, 宋小艺, 王忠宾, 杨建平, 徐坤. 2011. 嫁接与施肥对番茄产量及氮、磷、钾吸收利用效率的影响. 植物营养与肥料学报, 17 (1):131-136.
[34]	Zhang Z H, Cao B L, Gao S, Xu K. 2019. Grafting improves tomato drought tolerance through enhancing photosynthetic capacity and reducing ROS accumulation. Protoplasma, 256 (4):1013-1024. doi: 10.1007/s00709-019-01357-3 pmid: 30805718
[35]	Zheng Y J, Yang Z Q, Luo J, Zhang Y, Jiang N, Khattak W A. 2023. Transcriptome analysis of sugar and acid metabolism in young tomato fruits under high temperature and nitrogen fertilizer influence. Frontiers in Plant Science, 14: DOI:10.3389/fpls.2023.1197553.
[36]	Zhong Hai-xia, Zhang Fu-chun, Zhou Xiao-ming, Pan Ming-qi, Zhang Wen, Xie Hui, Han Shou-an, Ermek Caiksimu, Wu Xin-yu. 2018. Effects of different rootstocks on the accumulation of suger on Crimson Seedless grapes. Xinjiang Agricultural Sciences, 55 (6):1002-1010. (in Chinese) doi: 10.6048/j.issn.1001-4330.2018.06.003
	钟海霞, 张付春, 周晓明, 潘明启, 张雯, 谢辉, 韩守安, 艾尔买克·才卡斯木, 伍新宇. 2018. 不同砧木对无核葡萄果实糖分积累的影响. 新疆农业科学, 55 (6):1002-1010. doi: 10.6048/j.issn.1001-4330.2018.06.003
[37]	Zhou Jie, Shi Kai, Xia Xiaojian, Zhou Yanhong, Yu Jingquan. 2022. Vegetable cultivation technology in China:a sixty-year review and prospect. Acta Horticulturae Sinica, 49 (10):2131-2142. (in Chinese) doi: 10.16420/j.issn.0513-353x.2022-0858
	周杰, 师恺, 夏晓剑, 周艳虹, 喻景权. 2022. 中国蔬菜栽培科技60年回顾与展望. 园艺学报, 49 (10):2131-2142. doi: 10.16420/j.issn.0513-353x.2022-0858
[38]	Zhou Z, Yuan Y Q, Wang K T, Wang H J, Huang J Q, Yu H, Cui X. 2022. Rootstock-scion interactions affect fruit flavor in grafted tomato. Horticultural Plant Journal, 8 (4):499-510.

穗/砧 Scion/ Rootstock	果糖/（mg · g^-1FW） Fructose			葡萄糖/（mg · g^-1FW） Glucose			蔗糖/（mg · g^-1FW） Sucrose
穗/砧 Scion/ Rootstock	30 d	70 d	110 d	30 d	70 d	110 d	30 d	70 d	110 d
Dv/At	14.89 ± 1.12 a	14.37 ± 0.37 c	19.46 ± 0.65 c	20.71 ± 1.02 a	21.11 ± 0.84 b	22.58 ± 1.26 b	14.74 ± 0.42 a	8.04 ± 0.10 b	0.93 ± 0.11 e
Dv/Md	9.68 ± 0.26 c	16.23 ± 0.20 c	16.44 ± 0.85 d	15.61 ± 1.50 b	19.28 ± 0.20 c	25.37 ± 0.74 a	13.64 ± 0.51 b	9.48 ± 0.13 a	1.76 ± 0.12 d
Dv/Dv	11.72 ± 0.72 b	17.20 ± 0.15 b	20.72 ± 0.52 b	13.84 ± 0.20 c	24.27 ± 0.64 a	22.56 ± 0.87 b	10.53 ± 0.12 d	6.67 ± 0.20 c	2.76 ± 0.33 c
La/At	9.41 ± 0.21 c	12.45 ± 0.13 e	22.23 ± 1.10 a	15.87 ± 1.43 b	16.88 ± 0.82 d	21.92 ± 1.23 c	12.01 ± 0.38 c	5.86 ± 0.12 d	1.58 ± 0.14 d
La/Md	11.34 ± 0.50 b	15.42 ± 0.59 d	19.77 ± 0.65 c	16.42 ± 1.01 b	18.71 ± 0.64 c	17.53 ± 1.05 d	12.39 ± 0.16 c	6.90 ± 0.58 c	3.99 ± 0.23 b
La/La	7.34 ± 0.62 d	18.59 ± 0.48 a	19.83 ± 0.51 c	12.99 ± 0.70 c	21.75 ± 1.34 b	21.53 ± 1.25 c	10.86 ± 0.14 d	8.06 ± 0.31 b	4.83 ± 0.19 a
方差分析Variance analysis
接穗S	**	**	**	**	**	**	**	**	**
砧木R	**	**	**	**	**	ns	**	**	**
穗 × 砧 S × R	**	**	**	**	**	**	**	**	**

穗/砧 Scion/ Rootstock	果糖/（mg · g^-1FW） Fructose			葡萄糖/（mg · g^-1FW） Glucose			蔗糖/（mg · g^-1FW） Sucrose
穗/砧 Scion/ Rootstock	30 d	70 d	110 d	30 d	70 d	110 d	30 d	70 d	110 d
Dv/At	14.89 ± 1.12 a	14.37 ± 0.37 c	19.46 ± 0.65 c	20.71 ± 1.02 a	21.11 ± 0.84 b	22.58 ± 1.26 b	14.74 ± 0.42 a	8.04 ± 0.10 b	0.93 ± 0.11 e
Dv/Md	9.68 ± 0.26 c	16.23 ± 0.20 c	16.44 ± 0.85 d	15.61 ± 1.50 b	19.28 ± 0.20 c	25.37 ± 0.74 a	13.64 ± 0.51 b	9.48 ± 0.13 a	1.76 ± 0.12 d
Dv/Dv	11.72 ± 0.72 b	17.20 ± 0.15 b	20.72 ± 0.52 b	13.84 ± 0.20 c	24.27 ± 0.64 a	22.56 ± 0.87 b	10.53 ± 0.12 d	6.67 ± 0.20 c	2.76 ± 0.33 c
La/At	9.41 ± 0.21 c	12.45 ± 0.13 e	22.23 ± 1.10 a	15.87 ± 1.43 b	16.88 ± 0.82 d	21.92 ± 1.23 c	12.01 ± 0.38 c	5.86 ± 0.12 d	1.58 ± 0.14 d
La/Md	11.34 ± 0.50 b	15.42 ± 0.59 d	19.77 ± 0.65 c	16.42 ± 1.01 b	18.71 ± 0.64 c	17.53 ± 1.05 d	12.39 ± 0.16 c	6.90 ± 0.58 c	3.99 ± 0.23 b
La/La	7.34 ± 0.62 d	18.59 ± 0.48 a	19.83 ± 0.51 c	12.99 ± 0.70 c	21.75 ± 1.34 b	21.53 ± 1.25 c	10.86 ± 0.14 d	8.06 ± 0.31 b	4.83 ± 0.19 a
方差分析Variance analysis
接穗S	**	**	**	**	**	**	**	**	**
砧木R	**	**	**	**	**	ns	**	**	**
穗 × 砧 S × R	**	**	**	**	**	**	**	**	**

穗/砧 Scion/Rootstock	SS/（μmol · h^-1· g^-1FW）			SPS/（μmol · h^-1· g^-1FW）
穗/砧 Scion/Rootstock	30 d	70 d	110 d	30 d	70 d	110 d
Dv/At	7.52 ± 0.99 ab	12.01 ± 0.13 ab	12.94 ± 3.75 a	9.05 ± 0.25 a	11.83 ± 0.67 b	10.11 ± 0.43 bc
Dv/Md	8.87 ± 1.33 ab	10.07 ± 1.55 b	11.77 ± 0.98 a	9.35 ± 0.05 a	10.50 ± 0.17 c	11.37 ± 0.86 b
Dv/Dv	7.24 ± 0.02 b	11.73 ± 0.49 b	9.37 ± 0.33 b	7.53 ± 0.87 c	10.46 ± 1.39 c	8.48 ± 1.45 cd
La/At	8.93 ± 0.54 ab	9.82 ± 0.80 b	9.42 ± 1.36 b	8.89 ± 0.31 a	10.88 ± 0.10 bc	9.02 ± 0.72 cd
La/Md	7.98 ± 1.25 ab	11.53 ± 1.89 b	11.92 ± 0.20 a	8.13 ± 0.92 bc	10.48 ± 1.27 c	7.84 ± 0.89 d
La/La	9.32 ± 0.19 a	13.96 ± 1.85 a	11.03 ± 1.12 ab	8.73 ± 0.48 ab	13.79 ± 0.08 a	17.06 ± 1.34 a
方差分析Variance analysis
接穗Scion	ns	ns	ns	ns	ns	*
砧木Rootstock	ns	*	ns	ns	*	**
穗 × 砧S × R	*	*	**	*	**	**

穗/砧 Scion/Rootstock	SS/（μmol · h^-1· g^-1FW）			SPS/（μmol · h^-1· g^-1FW）
穗/砧 Scion/Rootstock	30 d	70 d	110 d	30 d	70 d	110 d
Dv/At	7.52 ± 0.99 ab	12.01 ± 0.13 ab	12.94 ± 3.75 a	9.05 ± 0.25 a	11.83 ± 0.67 b	10.11 ± 0.43 bc
Dv/Md	8.87 ± 1.33 ab	10.07 ± 1.55 b	11.77 ± 0.98 a	9.35 ± 0.05 a	10.50 ± 0.17 c	11.37 ± 0.86 b
Dv/Dv	7.24 ± 0.02 b	11.73 ± 0.49 b	9.37 ± 0.33 b	7.53 ± 0.87 c	10.46 ± 1.39 c	8.48 ± 1.45 cd
La/At	8.93 ± 0.54 ab	9.82 ± 0.80 b	9.42 ± 1.36 b	8.89 ± 0.31 a	10.88 ± 0.10 bc	9.02 ± 0.72 cd
La/Md	7.98 ± 1.25 ab	11.53 ± 1.89 b	11.92 ± 0.20 a	8.13 ± 0.92 bc	10.48 ± 1.27 c	7.84 ± 0.89 d
La/La	9.32 ± 0.19 a	13.96 ± 1.85 a	11.03 ± 1.12 ab	8.73 ± 0.48 ab	13.79 ± 0.08 a	17.06 ± 1.34 a
方差分析Variance analysis
接穗Scion	ns	ns	ns	ns	ns	*
砧木Rootstock	ns	*	ns	ns	*	**
穗 × 砧S × R	*	*	**	*	**	**

穗/砧 Scion/Rootstock	NI/（μmol · h^-1· g^{- 1}FW）			AI/（μmol · h^-1· g^-1FW）
穗/砧 Scion/Rootstock	30 d	70 d	110 d	30 d	70 d	110 d
Dv/At	29.78 ± 2.75 ab	30.35 ± 1.55 bc	49.35 ± 2.18 bc	25.29 ± 1.30 d	32.79 ± 2.88 c	264.52 ± 16.02 bc
Dv/Md	26.66 ± 3.00 b	32.42 ± 1.05 ab	63.87 ± 0.34 a	29.77 ± 2.29 c	34.22 ± 2.60 c	266.83 ± 13.71 bc
Dv/Dv	29.65 ± 1.39 ab	30.31 ± 2.57 bc	66.89 ± 3.17 a	32.06 ± 2.60 c	27.39 ± 2.60 d	295.63 ± 18.05 a
La/At	34.22 ± 2.88 a	27.04 ± 1.19 c	52.84 ± 3.25 b	41.63 ± 3.23 a	35.32 ± 0.91 bc	273.46 ± 18.11 ab
La/Md	26.51 ± 2.50 b	30.96 ± 2.31 b	45.04 ± 1.30 c	37.15 ± 0.66 b	39.82 ± 2.40 b	289.43 ± 3.92 ab
La/La	30.36 ± 2.57 ab	34.77 ± 0.86 a	52.08 ± 2.66 b	40.77 ± 2.93 ab	44.40 ± 2.00 a	247.23 ± 5.90 c
方差分析Variance analysis
接穗Scion	ns	ns	**	**	**	ns
砧木Rootstock	*	**	**	ns	ns	ns
穗 × 砧S × R	ns	**	**	*	**	*