黄绿叶突变体冀麦5265yg的光合生理特性分析

doi:10.3864/j.issn.0578-1752.2021.21.005

摘要/Abstract

摘要：

【目的】叶色突变体是研究叶绿素合成、叶绿体发育和光合作用的理想材料,探索小麦黄绿叶突变体的光合生理特性,旨在阐明其光合作用调控机理,为小麦黄绿叶突变体的进一步利用奠定基础。【方法】以野生型冀麦5265和突变体冀麦5265yg为试验材料,对叶色表型进行观察,采用分光光度计和试剂盒法测定色素含量和酶活性,并利用Li-6400便携式光合仪和PAM100叶绿素荧光仪进行光合气体交换参数和叶绿素荧光参数测定。【结果】表型观察和色素含量结果表明,突变体苗期叶片表现为黄绿色,抽穗后叶片逐渐转变为淡绿色。遮阴处理可以使叶片颜色部分复绿,但比野生型略浅,属于光诱导转绿型突变体。突变体叶绿素a和叶绿素b含量显著低于野生型,叶绿素a/b的比值升高,为典型的叶绿素缺乏型突变体;光响应曲线和CO₂响应曲线显示,突变体的表观量子效率（AQY）、光饱和点（LSP）、最大净光合速率（P_n-max）、光补偿点（LCP）、暗呼吸速率（Rd）、羧化效率（CE）和饱和CO₂浓度（I_-sat）显著高于野生型,说明突变体叶片的光合机构稳定,强光下光合速率更高;光合气体交换参数和叶绿素荧光动力学参数表明,突变体的净光合速率（P_n）、蒸腾速率（T_r）、气孔导度（G_s）、光化学量子效率（F_v/F_m）、实际光化学效率（ΦPSII）和光化学淬灭系数（qP）显著高于野生型,说明其具有较强的光能转化和CO₂固定能力;突变体的超氧化物歧化酶（SOD）和过氧化氢酶（CAT）活性均高于野生型,丙二醛（MDA）含量下降,可溶性糖和可溶性蛋白含量升高,说明抗氧化酶系统通过清除氧自由基降低了氧化损伤,突变体叶片细胞膜损害减轻,抗逆性增强;突变体的核酮糖-1,5-二磷酸羧化酶/加氧酶（Rubisco）活性显著低于野生型,磷酸烯醇式丙酮酸羧化酶（PEPC）活性显著高于野生型,推测C₄途径光合酶PEPC活性的升高可能是突变体具有较高净光合速率的原因之一。花后遮阴以及外源喷施抗坏血酸AsA和二硫苏糖醇DTT处理表明,突变体对光强变化更敏感、叶片内AsA含量及叶黄素循环效率更高。【结论】黄绿叶突变体冀麦5265yg叶片气孔导度明显改善、热耗散降低、C₄途径光合酶活性升高,是其光合速率提高的主要原因。该结果为小麦叶色突变体高光合特性的分子调控机制研究奠定基础。

关键词: 小麦, 黄绿叶突变体, 光合气体交换参数, 叶绿素荧光参数, 酶活性

Abstract:

【Objective】Leaf color mutants are ideal materials for studying chlorophyll synthesis, chloroplast development and photosynthesis. In order to clarify the regulation mechanism of photosynthesis and lay a foundation for the further utilization of wheat yellow-green leaf mutants, the photosynthetic physiological characteristics of wheat were studied.【Method】The wild type Jimai5265 and the mutant Jimai5265yg were used as test materials. The phenotype of leaf color was observed, the chlorophyll content and enzyme activity were measured by spectrophotometer and kit, respectively. The photosynthetic characteristics and chlorophyll fluorescence parameters were determined by the Li-6400 portable photosynthetic apparatus and PAM100 modulated chlorophyll fluorometer.【Result】The results of phenotypic observation and pigment content showed that the leaves of the mutant were yellow-green at seedling stage, and gradually changed to light green after heading stage. Leaf color of the mutant was partly recovered by shading treatment, but it was slightly lighter than the wild type, which indicated that it belonged to the mutants of light induced to promote greening. The content of chlorophyll a and b in the mutant leaves was significantly reduced, and the ratio of chlorophyll a to chlorophyll b was increased, indicating that Jimai5265yg was a typical chlorophyll deficient mutant. The light response curves and CO₂ response curve displayed that surface sight-seeing quantum efficiency (AQY), light saturation point (LSP), maximum net photosynthetic rate (P_n-max), light compensation point (LCP), dark respiration rate (Rd), spindle efficiency (CE) and saturated CO₂ concentration (I_-_sat) of the mutant was significantly higher than the wild type, indicating that the mutant had quite stable the photosynthetic mechanism and higher photosynthetic rate under the strong light; The photosynthetic gas exchange parameters and chlorophyll fluorescence kinetic parameters indicated that the net photosynthetic rate (P_n), transpiration rate (T_r), stomatal conductance (G_s), photochemical quantum efficiency (F_v/F_m), the actual photochemical efficiency (Φ PSII) and light chemical quenching coefficient (qP) of the mutant were significantly higher compared with the wild type, which showed that Jimai5265yg had the ability of strong light energy conversion and CO₂ fixation; The content of malondialdehyde (MDA) in mutant was decreased significantly, while the activity of superoxide dismutase (SOD) and catalase (CAT), as well as the content of soluble sugar and soluble protein, were significantly increased. The results indicated that the antioxidant enzyme system could reduce oxidative damage by scavenging oxygen free radicals. The damage degree of cell membrane in the mutant leaf was reduced and its stress resistance was enhanced. The activity of ribulose 1, 5-bisphosphate carboxylase/oxygenase (Rubisco) in mutant was significantly lower, while the activity of the phosphoenolpyruvate carboxylase (PEPC) was significantly higher than that of the wild type. It was speculated that the increased activity of the C₄ pathway photozyme PEPC might be the key factor for the higher net photosynthetic rate of the mutant. Post-flowering shading and exogenous spraying of Ascorbic acid and dithiothreitol DTT showed that the mutants were more sensitive to change of light intensity, and the content of AsA in leaves and the efficiency of xanthophyl cycle were higher.【Conclusion】Improvement of stomatal conductance, decrease of heat dissipation and increase of C₄ pathway photozyme activity in yellow green leaf mutant Jimai5265yg were the main reasons for the increase of photosynthetic rate. These results laid the foundation for the molecular regulation of high photosynthesis properties of wheat leaf mutants.

Key words: wheat, yellow-green leaf mutant, photosynthetic gas exchange parameters, chlorophyll fluorescence parameter, enzyme activity

郑伟, 师筝, 龙美, 廖允成. 黄绿叶突变体冀麦5265yg的光合生理特性分析[J]. 中国农业科学, 2021, 54(21): 4539-4551.

ZHENG Wei, SHI Zheng, LONG Mei, LIAO YunCheng. Photosynthetic and Physiological Characteristics Analysis of Yellow- Green Leaf Mutant in Wheat of Jimai5265yg[J]. Scientia Agricultura Sinica, 2021, 54(21): 4539-4551.

0
/ / 推荐

导出引用管理器 EndNote|Reference Manager|ProCite|BibTeX|RefWorks

链接本文: https://www.chinaagrisci.com/CN/10.3864/j.issn.0578-1752.2021.21.005

https://www.chinaagrisci.com/CN/Y2021/V54/I21/4539

图/表 12

图1

表1

图2

表2

表3

表4

图3

图4

图5

表5

图6

表6

参考文献 35

[36]	黄小辉, 冯大兰, 刘芸, 朱恒星, 陈道静, 耿养会. 模拟石漠化异质生境中桑树的生长和叶绿素荧光特性. 北京林业大学学报, 2016, 38(10):50-58.
	HUANG X H, FENG D L, LIU Y, ZHU H X, CHEN D J, GENG Y H. Growth and chlorophyll fluorescence characteristics of mulberry trees in simulated environment of heterogeneous habitats of a rocky desertification area. Journal of Beijing Forestry University, 2016, 38(10):50-58. (in Chinese)
[37]	AGARIE S, MIURA A, SUMIKURA R, TSUKAMOTO S, NOSE A, ARIMA S, MATAUOKA M, MIYAO-TOKUTOMI M. Overexpression of C4PEPC caused O-2-insensitive photosynthesis in transgenic rice plants. Plant Science, 2002, 162(2):257-265. doi: 10.1016/S0168-9452(01)00572-6
[38]	欧立军. 水稻叶色突变体的高光合特性. 作物学报, 2011, 37(10):1860-1867. doi: 10.3724/SP.J.1006.2011.01860
	OU L J. High photosynthetic efficiency of leaf colour mutant of rice (Oryza sativa L.). Acta Agronomica Sinica, 2011, 37(10):1860-1867. (in Chinese) doi: 10.3724/SP.J.1006.2011.01860
[39]	RUBAN A V. Identification of a mechanism of photoprotective energy dissipation in higher plants. Nature, 2007, 450(7169):575-578. doi: 10.1038/nature06262
[1]	CHEN H, CHENG Z J, MA X D, WU H, LIU Y L, ZHOU K N, CHEN Y L, MA W W, BI J C, ZHANG X, GUO X P, WANG J L, LEI C L, WU F Q, LIN Q B, LIU Y Q, LIU L L, JIANG L. A knockdown mutation of yellow-green leaf2 blocks chlorophyll biosynthesis in rice. Plant Cell Reports, 2013, 32(12):1855-1867. doi: 10.1007/s00299-013-1498-y
[2]	曹莉. 一个新的小麦黄化突变体研究[D]. 杨凌: 西北农林科技大学, 2007.
	CAO L. Characterization and genetics of a novel aurea mutant in wheat[D]. Yangling: Northwest A&F University, 2007. (in Chinese)
[3]	李宁. 小麦黄绿突变体特性研究与遗传分析[D]. 北京: 中国农业科学院, 2012.
	LI N. Characterization and genetic analysis of yellow green mutants in wheat[D]. Beijing: Chinese Academy of Agricultural Sciences, 2012. (in Chinese)
[4]	李倩倩. 小麦白斑突变体I30的特征特性及遗传分析[D]. 杨凌: 西北农林科技大学, 2017.
[40]	MASAHIRO I, NOZOMU U, FUMIHIKO S, TSUYOSHI E. Physiological functions of PsbS-dependent and PsbS-independent NPQ under naturally fluctuating light conditions. Plant and Cell Physiology, 2014, 55(7):1286-1295. doi: 10.1093/pcp/pcu069
[4]	LI Q Q. Characteristic and genetic analysis of common wheat mutant I30 with white stripe pattern[D]. Yangling: Northwest A&F University, 2017. (in Chinese)
[5]	CAMPBELL B W, MANI D, CURTIN S J, SLATTERY R A, MICHNO J M, ORT D R, SCHAUS P J, PALMER R G, ORF J H, STUPAR R M. Identical substitutions in magnesium chelatase paralogs result in chlorophyll-deficient soybean mutants. G3 Genes Genomes Genetics, 2015, 5(1):123-131.
[6]	孔可可, 许孟歌, 刘美凤, 孔杰杰, 盖钧镒, 赵团结. 大豆芽黄新突变体 vl-1的光合特性与基因定位. 核农学报, 2018, 32(5):840-847.
	KONG K K, XU M G, LIU M F, KONG J J, GAI J Y, ZHAO T J. Identification and fine mapping of a new virescent mutant vl-1 in soybean. Journal of Nuclear Agricultural Sciences, 2018, 32(5):840-847. (in Chinese)
[7]	钟世宜, 魏海忠, 王红红, 赵燕, 徐长利, 韩帅, 刘保申. 玉米白化突变体As-81647的鉴定及基因定位. 山东农业科学, 2013, 45(10):12-15.
	ZHONG S Y, WEI H Z, WANG H H, ZHAO Y, XU C L, HAN S, LIU B S. Identification and molecular mapping of an albino mutant gene As-81647 in maize (Zea mays L.). Shandong Agricultural Sciences, 2013, 45(10):12-15. (in Chinese)
[8]	江媛, 何筠, 范术丽, 俞嘉宁, 宋美珍. 棉花芽黄突变体10个叶绿体蛋白编码基因RNA编辑位点的测定及分析. 棉花学报, 2011, 23(1):3-9.
	JIANG Y, HE Y, FAN S L, YU J N, SONG M Z. The identification and analysis of RNA editing sites of 10 chloroplast protein-coding genes from virescent mutant of Gossypium hirsutum. Cotton Science, 2011, 23(1):3-9. (in Chinese)
[9]	宋明梅, 范术丽, 庞朝友, 魏恒玲, 喻树迅, 宋美珍. 棉花芽黄材料主要光合特性和农艺性状的研究. 棉花学报, 2015, 26(6):531-538.
	SONG M M, FAN S L, PANG C Y, WEI H L, YU S X, SONG M Z. Research on the main photosynthetic characteristics and agronomic traits in virescent cotton materials. Cotton Science, 2015, 26(6):531-538. (in Chinese)
[10]	孙捷音, 张年辉, 杜林方. 油菜叶绿素b减少突变体 Cr3529叶绿素生物合成的研究. 西北植物学报, 2007, 27(10):1962-1966.
	SUN J Y, ZHANG N H, DU L F. Chlorophyll biosynthesis in a chlorophyll b deficient oilseed rape mutant cr3529. Acta Botanica Boreali-Occidentalia Sinica, 2007, 27(10):1962-1966. (in Chinese)
[11]	殷家明, 杨惠娟, 彭柳, 黄梦珠, 唐章林, 李加纳, 李超. 甘蓝型油菜叶色黄化突变体Bn.el1研究. 西南大学学报(自然科学版), 2016, 38(5):1-6.
	YIN J M, YANG H J, PENG L, HUANG M Z, TANG Z L, LI J N, LI C. Preliminary research on the etiolation leaf-color mutant Bn. el1 in Brassica Napus. Journal of Southwest University (Natural Science Edition), 2016, 38(5):1-6. (in Chinese)
[12]	BRAUMANN I, STEIN N, HANSSO M. Reduced chlorophyll biosynthesis in heterozygous barley magnesium chelatase mutants. Plant Physiology and Biochemistry, 2014, 78:10-14. doi: 10.1016/j.plaphy.2014.02.004
[13]	QIN D D, DONG J, XU F C, GUO G G, GE S T, XU Q, XU Y X, LI M F. Characterization and fine mapping of a novel barley stage green-revertible albino gene (HvSGRA) by bulked segregant analysis based on SSR assay and specific length amplified fragment sequencing. BMC Genomics, 2015, 16(1):838. doi: 10.1186/s12864-015-2015-1
[14]	AWAN M A, KONZAK C, RUTGER J. Mutagenic effects of sodium azide in rice. Crop Science, 1980, 20:663-668. doi: 10.2135/cropsci1980.0011183X002000050030x
[15]	SHI J Q, WANG Y Q, GUO S, MA L, WANG Z W, ZHU X Y, SANG X C, LING Y H, WANG N, ZHAO F M, HE G H. Molecular mapping and candidate gene analysis of a yellow-green leaf 6 (ygl6) mutant in rice. Crop Science, 2015, 45(4):S41.
[16]	MA X Z, SUN X Q, LI C M, HUAN R, SUN C H, WANG Y, XIAO F L, WANG Q, CHEN P R, MA F R, ZHANG K, WANG P R, DENG X J. Map-based cloning and characterization of the novel yellow-green leaf gene ys83 in rice (Oryza sativa). Plant Physiology and Biochemistry, 2017, 111:1-9. doi: 10.1016/j.plaphy.2016.11.007
[17]	MEI J S, LI F F, LIU X R, HU G C, FU Y P, LIU W Z. Newly identified CSP41b gene localized in chloroplasts affects leaf color in rice. Plant Science, 2017, 256:39-45. doi: 10.1016/j.plantsci.2016.12.005
[18]	GUAN H Y, XU X B, HE C M, LIU C X, LIU Q, DONG R, LIU T S, WANG L M. Fine mapping and candidate gene analysis of the leaf-color gene ygl-1 in maize. PLoS ONE, 2016, 11(4):e0153962. doi: 10.1371/journal.pone.0153962
[19]	SHI D Y, ZHENG X, LI L, LIN W H, XIE W J, YANG J P, CHEN S J, JIN W W. Chlorophyll deficiency in the maize elongated mesocotyl2 mutant is caused by a defective heme oxygenase and delaying grana stacking. PLoS ONE, 2013, 8(11):e80107. doi: 10.1371/journal.pone.0080107
[20]	XING A Q, WILLIAMS M E, BOURETT T M, HU W N, HOU Z L, MEELEY R B, JAQUETH J, DAM T, LI B L. A pair of homoeolog ClpP5 genes underlies a virescent yellow-like mutant and its modifier in maize. Plant Journal, 2014, 79(2):192-205. doi: 10.1111/tpj.12568
[21]	LI W, TANG S, ZHANG S, SHAN J G, TANG C J, CHEN Q N, JIA G Q, HAN Y H, ZHI H, DIAO X M. Gene mapping and functional analysis of the novel leaf color gene SiYGL1 in foxtail millet (Setaria italica (L.) P. Beauv). Physiologia Plantarum, 2015, 157(1):24-37. doi: 10.1111/ppl.2016.157.issue-1
[22]	WANG Y K, HE Y J, YANG M, HE J B, XU P, SHAO M Q, CHU P, GUAN R Z. Fine mapping of a dominant gene conferring chlorophyll- deficiency in Brassica napus. Scientific Reports, 2016, 6:31419. doi: 10.1038/srep31419
[23]	WANG R, YANG F, ZHANG X Q, WU D X, TAN C, WESTCOTT S, BROUGHTON S, LI C D, ZHANG W Y, XU Y H. Characterization of a thermo-inducible chlorophyll-deficient mutant in barley. Plant Science, 2017, 14(8):1936.
[24]	秦丹丹, 李梅芳, 许甫超, 徐晴, 葛双桃, 董静. 大麦黄绿叶色突变体ygl的农艺性状及其调控基因初步定位. 麦类作物学报, 2019, 39(6):653-658.
	QIN D D, LI M F, XU B C, XU Q, GE S T, DONG J. Analysis of agronomic characters and preliminary mapping of regulatory genes of a barley yellow-green leaf mutant ygl. Journal of Triticeae Crops, 2019, 39(6):653-658. (in Chinese)
[25]	ZHANG L L, LIU C, AN X Y, WU H Y, FENG Y, WANG H, SUN D J. Identification and genetic mapping of a novel incompletely dominant yellow leaf color gene, Y1718, on chromosome 2BS in wheat. Euphytica, 2017, 213(7):141. doi: 10.1007/s10681-017-1894-4
[26]	WU H Y, SHI N R, AN X Y, LIU C, FU H F, CAO L, FENG Y, SUN D J, ZHANG L L. Candidate genes for yellow leaf color in common wheat (Triticum aestivum L.) and major related metabolic pathways according to transcriptome profiling. International Journal of Molecular Sciences, 2018, 19(6):1594. doi: 10.3390/ijms19061594
[27]	茹广欣, 刘小囡, 朱秀红, 张龙冲, 王鋆瑞, 周霜晴. 泡桐黄化突变体生理特性分析. 南京林业大学学报(自然科学版), 2017, 41(4):181-185.
	RU G X, LIU X N, ZHU X H, ZHANG L C, WANG J R, ZHOU S Q. Physiological characteristic analysis of etiolation mutant in Paulownia fortnnei. Journal of Nanjing Forestry University (Natural Sciences Edition), 2017, 41(4):181-185. (in Chinese)
[28]	杨小苗, 吴新亮, 刘玉凤, 李天来, 齐明芳. 一个番茄EMS叶色黄化突变体的叶绿素含量及光合作用. 应用生态学报, 2018, 29(6):1983-1989.
	YANG X M, WU X L, LIU Y F, LI T L, QI M F. Analysis of chlorophyll and photosynthesis of a tomato chlorophyll-deficient mutant induced by EMS. Chinese Journal of Applied Ecology, 2018, 29(6):1983-1989. (in Chinese)
[29]	胡亮亮, 赵子瑶, 张海强, 陈菲帆, 张朝文, 戎福喜, 陈鹏, 李玉红. 一个新的黄瓜叶色突变体的光合特性分析. 西北农业学报, 2018, 27(11):1622-1628.
	HU L L, ZHAO Z Y, ZHANG H Q, CHEN F F, ZHANG C W, WU F X, CHEN P, LI Y H. Photosyntheic characteristic analysis of new leaf color mutant in cucumber. Acta Agriculturae Boreali-Occidentalia Sinica, 2018, 27(11):1622-1628. (in Chinese)
[30]	曹莉, 王辉, 孙道杰, 冯毅, 李学军, 闵东红. 小麦黄化突变体类囊体蛋白组分及叶绿素的合成特性. 麦类作物学报, 2010, 30(4):638-643.
	CAO L, WANG H, SUN D J, FENG Y, LI X J, MIN D H. Chloroplast thylakoid protein composition and characteristics of chlorophyll biosynthesis in a novel aurea mutant of wheat. Journal of Triticeae Crops, 2010, 30(4):638-643. (in Chinese)
[31]	DAI X B, XU X M, LU W. Photoinhibition characterristics of a low chlorophyll b mutant of high yield rice. Photosynthetica, 2003, 41:57-60. doi: 10.1023/A:1025804327776
[32]	ZHOU X S, SHEN S Q, WU D X, SUN J W, SHU Q Y. Introduction of a xantha mutation for testing and increasing varietal purity in hybrid rice. Field Crops Research, 2006, 96(1):71-79. doi: 10.1016/j.fcr.2005.05.008
[33]	DENG X J, ZHANG H Q, WANG Y, HE F, LIU J L, XIAO X, SHU Z F, LI W, WANG G H, WANG G L. Mapped clone and functional analysis of leaf-color gene Ygl7 in a rice hybrid (Oryza sativa L. ssp. indica). PLoS ONE, 2014, 9(6):e99564. doi: 10.1371/journal.pone.0099564
[34]	LICHTENTHALER H K. Chlorophylls and carotenoids: Pigments of photosynthetic biomembranes. Methods Enzymol, 1987, 148:350-382.
[35]	叶子飘. 光合作用对光和CO₂响应模型的研究进展. 植物生态学报, 2010, 34(6):727-740 doi: 10.3773/j.issn.1005-264x.2010.06.012
	YE Z P. A review on modeling of responses of photosynthesis to light and CO₂. Chinese Journal of Plant Ecology, 2010, 34(6):727-740. (in Chinese) doi: 10.3773/j.issn.1005-264x.2010.06.012

材料 Material	叶绿素 a Chl a	叶绿素 b Chl b	类胡萝卜素 Car	叶绿素a/b Chl (a/b)	总叶绿素 Total Chl	胡萝卜素/叶绿素 Car/Chl ratio
冀麦5265 Jimai5265	1.06±0.20a	0.31±0.05a	0.23±0.03a	3.44±0.08b	1.36±0.25a	0.17±0.01b
冀麦5265yg Jimai5265yg	0.65±0.07b	0.13±0.01b	0.20±0.02a	5.05±0.36a	0.78±0.09b	0.25±0.06a

特征参数值 Parameter	材料 Material
特征参数值 Parameter	冀麦5265 Jimai5265	冀麦5265yg Jimai5265yg
表观量子效率 AQY (µmol·µmol^-1)	0.0669±0.009b	0.0719±0.006a
光饱和点 LSP (µmol·m^-2·s^-1)	1619±35.69b	1873±56.32a
光补偿点 LCP (µmol·m^-2·s^-1)	37.65±4.19b	56.84±5.63a
最大净光合速率 P_n-max (µmol·m^-2·s^-1)	29.47±1.34b	45.96±2.28a
暗呼吸速率 Rd (µmol·m^-2·s^-1)	2.44±0.74b	3.82±0.59a
羧化效率 CE (mol·m^-2·s^-1)	0.17±0.03b	0.25±0.04a
光合能力 A_n-max(µmol·m^-2·s^-1)	41.16±3.12b	50.85±6.24a
饱和CO₂浓度 I_-sat (µmol·mol^-1)	610.93±53b	890.14±74a
CO₂补偿点 Γ (µmol·mol^-1)	56.51±3.18a	51.66±2.12b
光呼吸速率 Rp (mol·m^-2·s^-1)	8.74±0.09a	11.40±0.24b

生育时期 Growth stage	材料 Material	净光合速率 Pn (µmol·m^-2·s^-1)	气孔导度 Gs (mmol·m^-2·s^-1)	胞间CO₂浓度 Ci (µmolmol^-1)	蒸腾速率 Tr (mmol·m^-2·s^-1)
拔节期 Jointing stage	冀麦5265 Jimai5265	10.67±0.99	0.13±0.02	245.07±43.51	3.65±0.68
拔节期 Jointing stage	冀麦5265yg Jimai5265yg	13.82±1.81*	0.14±0.06	249.93±38.02	4.00±1.18
开花期 Flowering stage	冀麦5265 Jimai5265	20.95±1.37	0.38±0.06	286.92±17.33	7.59±0.62
开花期 Flowering stage	冀麦5265yg Jimai5265yg	25.63±1.02*	0.66±0.13*	306.59±14.86*	8.54±0.43*
灌浆期 Filling stage	冀麦5265 Jimai5265	15.89±0.77	0.27±0.05	98.25±12.67	6.12±0.69
灌浆期 Filling stage	冀麦5265yg Jimai5265yg	16.93±1.05*	0.32±0.05	109.40±6.88	7.34±0.92*

参数 Parameter	拔节期 Jointing stage		开花期Flowering stage		灌浆期Filling stage
参数 Parameter	冀麦5265 Jimai5265	冀麦5265yg Jimai5265yg	冀麦5265 Jimai5265	冀麦5265yg Jimai5265yg	冀麦5265 Jimai5265	冀麦5265yg Jimai5265yg
F₀	0.331±0.009	0.201±0.014**	0.350±0.015	0.315±0.016*	0.394±0.010	0.335±0.013*
F_0'	0.233±0.005	0.186±0.005**	0.318±0.012	0.291±0.017	0.380±0.011	0.347±0.016
F_m'	0.477±0.012	0.633±0.039**	1.305±0.056	1.293±0.141	1.514±0.065	1.362±0.132
F_v/F_m	0.729±0.009	0.758±0.014*	0.831±0.007	0.841±0.010*	0.817±0.008	0.817±0.006
Φ_qP	0.306±0.030	0.681±0.040**	0.756±0.010	0.775±0.015*	0.748±0.011	0.744±0.014
Φ_PSII	0.322±0.030	0.486±0.039**	0.572±0.041	0.611±0.029	0.578±0.027	0.616±0.008*
qP	0.630±0.073	0.690±0.054	0.773±0.054	0.789±0.040	0.772±0.028	0.827±0.019
NPQ	1.566±0.128	0.323±0.206**	0.601±0.089	0.533±0.088	0.151±0.066	0.070±0.033**
qL	0.456±0.083	0.400±0.067	0.459±0.075	0.462±0.064	0.461±0.034	0.550±0.043

材料 Material	果糖 Fructose	葡萄糖 Glucose	蔗糖 Sucrose	松三糖 Melezitose
冀麦5265 Jimai5265	12.942±3.564a	24.073±4.644a	15.737±9.366b	0.032±0.022b
冀麦5265yg Jimai5265yg	17.442±5.497a	28.749±7.785a	60.67±15.500a	0.091±0.045a