杂交中稻开花期高温对结实率影响及其与组合间库源结构和开花习性的关系

引用本文

徐富贤, 张林, 熊洪, 朱永川, 刘茂, 蒋鹏, 郭晓艺. . 杂交中稻开花期高温对结实率影响及其与组合间库源结构和开花习性的关系. 作物学报, 2015, 41(6): 946-955
XU Fu-Xian, ZHANG Lin, XIONG Hong, ZHU Yong-Chuan, LIU Mao, JIANG Peng, GUO Xiao-Yi. . Effects of High Temperature during Flowering Period on Seed Setting Rate and Its Relationship with Sink to Source Ratios and Flowering Habit of Mid-season Hybrid Rice. ACTA AGRONOMICA SINICA, 2015, 41(6): 946-955 复制到剪切板

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杂交中稻开花期高温对结实率影响及其与组合间库源结构和开花习性的关系

徐富贤^1,^2,^*, 张林^1,^2,^*, 熊洪^1,², 朱永川¹, 刘茂¹, 蒋鹏¹, 郭晓艺¹

¹四川省农业科学院水稻高粱研究所 / 农业部西南水稻生物学与遗传育种重点实验室, 四川德阳618000

²国家水稻改良中心四川泸州分中心, 四川泸州646100

^* 通讯作者(Corresponding author): 徐富贤, E-mail: Xu6501@163.com, Tel: 18090167012 ^*同等贡献(Contributed equally to this work)

收稿日期:2014-11-19 接受日期:2015-04-02 网络出版日期:2015-04-21

基金:本研究由国家公益性行业(农业)科研专项(201203029), 国家粮食科技丰产工程项目(2011BAD16BO5-1), 四川省财政基因工程项目(2013LWJJ-009), 西南水稻创新体系建设项目和四川省财政推广项目资助

摘要

2011—2013年, 以50个杂交中稻组合为材料, 通过大田分期播种与智能人工气候室控温, 以使开花期遇高温伤害, 研究了杂交组合开花期的耐高温特性与库源性状及开花习性的关系。结果表明, 杂交组合间耐高温能力差异显著, 鉴定出了开花期常温下结实率和耐高温指数均较高的组合10个, 即绵香优576、蓉稻优415、香绿优727、绵优5240、川谷优202、川优6203、冈优169、内香优5828、川谷优7329、江优126。开花期耐高温能力与开花当时植株光合物质的供给水平有关, 有效穗数越少的组合的穗粒数和粒叶比越高, 即单位颖花的光合源占有量越少, 以致开花期耐高温指数与粒叶比呈显著负相关; 9:30—11:00时段开花比例和叶绿素含量分别与耐高温指数呈显著正相关。选用穗数型、花时早而集中的杂交组合和提高开花期叶绿素含量, 是增强杂交水稻开花期耐高温能力的重要途经。

关键词: 杂交中稻; 开花期; 耐高温指数; 库源结构; 开花习性

Effects of High Temperature during Flowering Period on Seed Setting Rate and Its Relationship with Sink to Source Ratios and Flowering Habit of Mid-season Hybrid Rice

XU Fu-Xian^1,^2,^*, ZHANG Lin^1,^2,^*, XIONG Hong^1,², ZHU Yong-Chuan¹, LIU Mao¹, JIANG Peng¹, GUO Xiao-Yi¹

¹ Rice and Sorghum Research Institute, Sichuan Academy of Agricultural Sciences / Key Laboratory of Southwest Rice Biology and Genetic Breeding, Ministry of Agriculture, Deyang 618000, China

² Luzhou Branch of National Rice Improvement Center, Luzhou 646100, China

Abstract

From 2011 to 2013, under the condition of high temperature injury during heading stage designed by applying different sowing dates in the field and controlling the temperature in intelligent artificial climate chamber, the relationship high temperature resistance index (HTRI) with sink to source ratio and flowering habit in mid-season hybrid rice was studied with 50 hybrid combinations as tested materials. By using variance, correlation and regression analyses, the results showed that there were significant differences in HTRI among hybrid combinations. Ten of the combinations were tested to show a higher levels of seed setting rate and HIRI during flowering stage, including Mianxiangyou 576, Rongdaoyou 415, Xianglüyou 727, Mianyou 5240, Chuanguyou 202, Chuanyou 6203, Gangyou 169, Neixiangyou 5828, Chuanguyou 7329, and Jiangyou 126. There existed a significant relation between HTRI and carbohydrate supply level during flowering stage. The smaller the number of effective panicles of hybrid combinations was, the higher the spikelets per panicle and grain-leaf ratio would be, namely, the less the occupancy of photosynthetic source per spikelets would be. As a consequence, significantly negative correlation were observed between HTRI and grain-leaf ratio during flowering stage. The flowering ratio from 9:30 to 11:00 and chlorophyll content of leaf had a significantly positive correlation with HTRI respectively. Choosing the hybrid combinations with a larger number of effective panicles and early intensive anthesis and raising the level of chlorophyll content of leaf are the vital ways to enhance HTRI during heading stage.

Keyword: Mid-season hybrid rice; Flowering stage; High temperature resistance index; Sink to source ratio; Flowering habit

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引言

全球气候变暧已成为一个不争的事实^{[1, 2]}, 它将对整个陆地生态系统的碳循环产生重大影响^[1]。水稻是我国南方的主要粮食作物之一, 在全球气候变暖的大环境下, 极端天气频繁发生, 高温已经成为制约水稻产量和品质的主要因素^{[3, 4]}。特别是抽穗开花期的极端高温造成水稻结实率显著下降而大幅度减产^{[5, 6]}。因此, 开展水稻品种开花期耐高温机理及其缓解技术研究是近年国内外的热点课题^{[4, 5, 7, 8]}。

品种选择是抵御高温危害的重要措施^{[9, 10]}。因此, 国内外就水稻在高温胁迫下叶片的光合特性及叶绿素含量、可溶性糖、可溶性蛋白、游离脯氨酸、热稳定蛋白、膜透性、MDA含量和光合作用的关键酶Rubisco、活化酶RuBP、羧化酶活性^{[11, 12, 13, 14, 15, 16]}, 花粉的活力、萌发率、可育率、受精率、结实率及花器官发育^{[17, 18, 19, 20]}, 籽粒的蔗糖、果糖和葡萄糖含量的动态变化以及蔗糖合酶、液泡型转化酶和细胞壁结合型转化酶活性等^[21]的生理生化响应, 开展了广泛而深入的研究。为破解水稻品种间耐高温差异性的机理提供了科学依据。然而, 水稻遗传育种是一门实践性很强的学科, 利用前述研究成果难以直接应用于耐高温品种的选育工作, 在水稻品种选育过程中, 十分需要明确与耐高温有关的植株形态特征, 作为鉴定开花期耐高温能力的选择依据, 而该方面的研究文献极少。为此, 本文以多个杂交水稻组合为材料, 开展了杂交水稻开花期高温对结实率的影响与组合库源性状及开花习性的关系研究, 以期为开花期耐高温品种的选育提供理论依据。

1 材料与方法

试验在四川省农业科学院水稻高粱研究所泸县实验基地的冬水田及智能人工气候室进行, 田间试验土壤质地均匀, 中上等肥力。

1.1 自然高温对结实率影响与库源性状和开花习性的关系田间试验

2011年以30个杂交中稻组合(表1)为材料, 分别于3月5日、4月5日、4月20日、6月1日、6月25日播种, 2013年以40个杂交中稻组合为材料, 分别于3月5日、3月25日、4月15日、5月5日、6月15日播种(为终有一个播期的杂交组合在抽穗期能遇高温)。湿润育秧, 4.5叶期移栽。大田按30 cm × 20 cm规格每穴栽双株, 本田施氮150 kg hm^-2 (底肥∶ 蘖肥∶ 穗肥=5∶ 3∶ 2), 磷、钾肥按N∶ P₂O₅∶ K₂O = 1.0∶ 0.5∶ 0.8作底肥一次施用。2011年每期各杂交组合移栽120穴, 重复3次, 裂区设计; 2013年每个组合栽100穴。以播期为主区, 杂交组合为副区。

2011年考查各杂交组合的始穗期、齐穗期, 并分别统计各播期的杂交组合抽穗期(齐穗前5 d至齐穗后3 d)的日最低气温、日最高气温和日平均气温。气温资料来源于距试验田1000 m、海拔相差10 m左右的泸县气象局气象观测站。本田苗期采用常规方法考查最高苗数, 齐穗期采用干物重法考查3月5日播种的各杂交组合的粒叶比^[22], 成熟期各播期所有组合的每次重复均取样5穴, 考查穗部性状, 抽穗期的耐高温指数=高温下结实率/常温下结实率× 100%。

2013年观察到对水稻开花受精具有明显伤害的高温(日最高温度达35℃)^[6]稳定期在7月11日下午。此高温期间由于在5个播种期中只有3月25日播种的绝大多数杂交组合正处于抽穗期。因此, 在3月25日播种的40个杂交组合中选择20个(内5优306、川优6203、蓉18优447、冈优169、宜香优800、冈比优99、金冈优983、内5优317、川农优华占、内香优5828、川谷优7329、蓉优22、冈优618、T优109、科优21、江优126、中优849、金优T16、奇优801、花香优1号)同期开始抽穗、生长整齐的稻穗挂牌标记, 每个杂交组合每次重复挂牌标记6~8穗, 重复3次。于7月12日至15日高温期间, 分别统计9:30以前、9:30— 11:00、11:00— 12:30和12:30以后的开花数, 并从7:30至15:30, 每小时记录1次穗层气温。齐穗期将3月5日播期(常温下抽穗)中与3月25日(高温下抽穗)挂牌标记相同的20个组合, 各取样5穴, 3次重复, 考查粒叶比^[22]、顶部3叶叶绿素含量(SPAD值)^[23]和单茎干物重; 成熟期将3月5日播种与3月25日挂牌标记相同的20个组合, 各取样5穴, 3次重复, 从3月25日播种期中选定的20个组合挂牌标记定穗的穗子全部取样, 考查穗部性状, 并计算各杂交组合抽穗期的耐高温指数。

1.2 智能人工气候室试验

试验在温度精确度± 0.5℃的智能人工气候室(北京易盛泰和科技有限公司制)进行。采用钵栽试验, 取稻田干土, 经晒场整细混均后等量装入钵内。2012年以杂交中稻组合绵优5240为材料, 3月8日播种, 地膜培育中苗秧, 4.5叶移栽至钵盆。氮按每钵1.5 g施用, 其中底肥50%、蘖肥20%、穗肥30%, 磷、钾肥按N∶ P₂O₅∶ K₂O = 1.0∶ 0.5∶ 1.0比例作底肥一次性施入。于抽穗前剪叶处理, 分别设剪去绿叶面积的1/4 (沿叶脉剪掉半叶上部的1/2)、1/3 (剪掉整叶上部的1/3)、1/2 (沿叶脉剪掉一半叶)、3/4 (先沿叶脉剪掉一半叶, 再将剩下的半叶剪掉上部的1/2)和不剪叶5个处理。2013年以杂交中稻组合冈优169为材料, 3月5日播种, 地膜培育中苗秧, 五叶期移栽至盆钵。设6个施氮水平(0、0.5、1.0、1.5、2.0、2.5 g 钵^-1), 其中底肥60%、蘖肥20%、穗肥20%, 磷、钾肥按N∶ P₂O₅∶ K₂O = 1.0∶ 0.5∶ 0.5比例作底肥一次性施入。

各处理栽13~15钵, 每钵栽3穴, 每穴栽双株。始穗期每处理选择生长基本一致的10钵, 将其中5钵移至人工气候室高温处理5 d (相对湿度85%, 温度为7:00— 9:00 34.5℃, 9:00— 14:00 38.0℃, 14:00— 16:00 34.5℃, 16:00— 7:00 31.0℃), 另5钵置田间常温下, 高温处理期间保持盆钵内浅水层。高温处理结束后将盆钵移至田间正常管理。于成熟期分别收获, 考查各处理结实率和耐高温指数, 其中2013年试验在移至人工气候室高温处理前测顶部3片叶的叶绿素含量(SPAD值)^[23]和粒叶比^[22]。

以上试验数据的方差、相关及回归分析由DPS数据处理系统和Microsoft Excel操作系统完成。

2 结果与分析

2.1 群体条件下高温对结实率影响及与组合间库源性状关系

从表1可见, 5个播种期下抽穗期的温度状况明显不同, 其中只有6月1日播种的遇到35℃以上的高温(较常年明显推迟), 6月25日遇到22℃以下的低温。这两期的结实率均比前3期显著下降。前3期抽穗期间气温正常, 结实率差异不显著, 但以3月5日播期略高。因此, 以3月5日播期的结实率作为常温对照。再从6月1日播种的30个杂交组合抽穗期9 d的气温(表2)看, 除宜香优2079、川香优858、宜香优4106三个组合日最高温平均接近35℃以外, 其余27个组合均在35℃以上高温下抽穗, 但各杂交组合间抽穗期的绝对温度条件有一定差异, 造成一定的结实差异。30个杂交组合分别在常温和高温下开花的结实率及其耐高温指数差异极显著, 其中在常温下结实率较高( > 80%), 耐高温指数达85%以上的5个杂交组合有绵香优576、蓉稻优415、香绿优727、绵优5240和川谷优202。

30个杂交组合间的穗粒结构及库源结构均达极显著差异(方差分析F值2.06^{* *}~40.48^{* *}), 相关分析结果显示, 杂交组合间的最高苗、有效穗、穗粒数、粒叶比4个性状与耐高温指数有显著相关性(表3), 其中2011年有效穗(x₂)和粒叶比(x₈)对耐高温指数的偏相关达显著或极显著水平(表4)。究其原因, 有效穗数越低的组合, 其穗粒数越高(图1), 穗粒数越高组合的粒叶比越高即单位颖花的光合源占有量越少(图2), 以致抽穗期耐高温指数与粒叶比呈极显著负相关(图3), 2011和2013两年结果表现一致。表明抽穗开花期遇高温情况下有充足的光合物质供应, 有利于减轻对颖花受精结实的危害。

2.2 定穗条件下高温对结实率影响及与组合间库源结构和开花习性的关系

从定穗观察20个组合开花习性的7月12日至15日连续4 d气温状况看, 均出现了35℃以上的日最高温, 而且均出现在12:30以后(表5)。在4 d时间中所有20个组合开花均结束。表6显示, 定穗条件下杂交组合间的结实率、耐高温指数、不同时段开花比例、库源结构的差异显著或极显著(方差分析F值2.24^*~23.34^{* *}), 由于该年度试验抽穗期遇高温仅5 d, 定穗开花期结束后的7月16日至20日为多云转晴或小雨, 日最高气温25.0~34.5℃, 其高温强度明显比前试验弱。因此, 20个参试组合的平均耐高温指数高达94.55%, 比前试验30个组合平均耐高温指数78.54%高16.01个百分点。在本试验中同时满足常温下开花结实率85%以上、耐高温指数95%以上的5个组合有川优6203、冈优169、内香优5828、川谷优7329和江优126。

从相关分析结果看, 9:30— 11:00开花越多耐高温指数越高, 而11:00— 12:30和12:30以后开花越多则耐高温指数越低, 耐高温指数分别与粒叶比和穗粒数呈显著或极显著负相关(表6); 其中9:30— 11:00开花比例(z₂)和穗粒数(z₆)的偏相关达显著或极显著水平(表4)。表明9:30— 11:00开花比例高和穗粒数少的组合耐高温能力较强。分析其原因, 开花期高温伤害的敏感期在开花当时, 对受精影响最大, 花后1 h则无明显影响^[24], 而本试验35 ℃以上的日最高温出现在12:30以后, 9:30— 11:00开花比例高的组合因避开了高温伤害敏感期而减轻了伤害; 穗粒数高影响耐高温指数应与较低的光合源供给水平有关(图2和图3), 与前试验结果表现一致。

表1 30个杂交组合不同播期下抽穗期的温度状况与结实率(2011) Table 1 Temperature and seed setting rate at heading stage of 30 hybrid combinations with different sowing dates (2011)

表2 30个杂交组合抽穗期的高温状况及耐高温指数(2011) Table 2 High temperature at heading stage and high temperature resistance index of 30 hybrid combinations (2011)

杂交组合 Hybrid combination	始穗期 BH (M/D)	齐穗期 FH (M/D)	抽穗期HS (始穗-齐穗后3 d)			结实率 Seed setting rate (%)		耐高温指数 HTRI (%)
杂交组合 Hybrid combination	始穗期 BH (M/D)	齐穗期 FH (M/D)	MTD (℃)	HTD (℃)	LTD (℃)	高温期 HTS	常温期 NTS	耐高温指数 HTRI (%)
G优802 G you 802	8/25	8/29	29.1	36.5	23.4	36.8 l	74.7 h	49.3 m
绵香优576 Mianxiangyou 576	8/21	8/25	29.4	35.3	24.7	72.9 bc	83.1 bcdef	87.7 bcd
蓉18优188 Rong 18 you 188	8/20	8/24	30.1	35.9	25.5	63.6 fgh	78.7 defgh	80.8 defgh
宜香优7808 Yixiangyou 7808	8/23	8/27	28.6	35.0	23.5	55.1 ij	76.8 efgh	71.8 jk
宜香优2079 Yixiangyou 2079	8/24	8/27	28.2	35.0	23.1	54.9 ij	81.4 bcdefgh	67.4 kl
D优6511 D you 6511	8/25	8/29	29.1	36.5	23.4	63.9 efgh	84.7 abcd	75.5 ij
内2优6号 Nei 2 you 6	8/24	8/28	28.5	35.5	23.4	59.2 gh	75.6 gh	78.3 fghij
川香优858 Chuanxiangyou 858	8/25	8/28	29.2	34.9	24.8	70.9 bcde	79.4 defgh	89.3 abc
川香优3203 Chuanxiangyou 3203	8/25	8/29	29.1	36.5	23.4	60.9 fgh	79.8 cdefgh	76.3 ghij
内5优39 Nei 5 you 39	8/21	8/25	29.4	35.3	24.7	77.0 ab	90.7 a	84.9 bcdef
蓉稻优415 Rongdaoyou 415	8/22	8/25	29.3	35.3	24.6	72.6 bcd	82.5 bcdefg	87.9 bcd
宜香优2168 Yixiangyou 2168	8/20	8/24	30.1	35.9	25.5	62.3 fgh	81.5 bcdefgh	76.4 ghij
泰优99 Taiyou 99	8/25	8/29	29.1	36.5	23.4	72.1 bcd	78.4 defgh	91.9 ab
内5优317 Nei 5 you 317	8/22	8/26	28.9	35.1	24.0	65.5 defg	83.5 bcde	78.5 fghij
川作6优177 Chuanzuo 6 you 177	8/19	8/23	30.9	36.8	26.3	67.7 cdef	81.0 bcdefgh	83.6 defgh
内香优18 Neixiangyou 18	8/25	8/29	29.1	36.5	23.4	48.2 k	77.0 efgh	62.5 l
香绿优727 Xianglü you 727	8/22	8/26	28.9	35.1	24.0	72.5 bcd	81.1 bcdefgh	89.4 abc
宜香优4245 Yixiangyou 4245	8/22	8/26	28.9	35.1	24.0	66.4 cdefg	76.8 efgh	86.4 bcde
内香优8156 Neixiangyou 8156	8/20	8/24	30.1	35.9	25.5	76.0 ab	91.0 a	83.6 cdefgh
冈香优707 Gangxiangyou 707	8/19	8/23	30.3	36.1	25.8	48.0 k	87.6 ab	54.84m
绵优5240 Mianyou 5240	8/20	8/24	30.1	35.9	25.5	76.8 ab	84.7 abcd	90.7 abc
内香优2128 Neixiangyou 2128	8/22	8/26	28.9	35.1	24.0	63.8 efgh	76.1 fgh	83.8 cdefg
内香优2550 Neixiangyou 2550	8/22	8/27	29.1	35.5	24.1	68.1 cdef	75.7 gh	90.0 abc
内5优5399 Nei 5 you 5399	8/20	8/24	30.1	35.9	25.5	61.6 fgh	82.0 bcdefg	75.1 ij
宜香优7633 Yixiangyou 7633	8/22	8/26	28.9	35.1	24.0	51.8 jk	78.8 defgh	65.7 kl
宜香优4106 Yixiangyou 4106	8/22	8/25	28.8	34.8	24.0	59.2 gh	77.6 defgh	76.3 ghij
川谷优202 Chuanguyou 202	8/22	8/26	28.9	35.1	24.0	80.2 a	83.3 bcdef	96.4 a
乐丰优329 Lefengyou 329	8/22	8/26	28.9	35.1	24.0	66.4 cdefg	87.5 ab	75.9 hij
宜香优1108 Yixiangyou 1108	8/20	8/24	30.1	35.9	25.5	65.4 defg	82.0 bcdefg	79.8 efgh
川香优727 Chuanxiangyou 727	8/23	8/26	29.5	35.3	24.9	57.5 hij	86.8 abc	66.2 kl
F值 F-value						20.6^{* *}	4.4^{* *}	22.6^{* *}

BH: heading beginning; FH: full heading; HS: heading stage (from BH to 3 days after FH); MTD: mean daily temperature; HTD: the highest daily temperature; LTD: the lowest daily temperature; HTS: high temperature stage; HTRI: high temperature resistance index; NTS: normal temperature stage. Values within a column followed by different letters are significantly different at the 0.05 probability level.

同一列中数据后跟不同字母表示在0.05水平差异显著。

表2 30个杂交组合抽穗期的高温状况及耐高温指数(2011) Table 2 High temperature at heading stage and high temperature resistance index of 30 hybrid combinations (2011)

表3 30个杂交组合常温下抽穗期库源性状与耐高温指数的关系(2011) Table 3 Relationship between the sink-source characteristic and high temperature resistance index during heading stage at normal temperature in 30 hybrid combinations (2011)

表4 抽穗期耐高温指数(y)与植株性状(x或z)的多元回归分析 Table 4 Multiple regression analysis between high temperature resistance index (y) at heading stage and characteristic of above ground plants (x or z)

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	图1 杂交组合穗粒数与有效穗数关系(2011)Fig. 1 Relationship between spikelets per panicle and effective panicles of hybrid combinations (2011)

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	图2 杂交组合间粒叶比与穗粒数关系Fig. 2 Relationship between grain-leaf ratio and spikelets per panicle of hybrid combinations

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	图3 抽穗期耐高温指数与粒叶比关系Fig. 3 Relationship between high temperature resistance index (HTRI) and grain-leaf ratio at heading stage

表5 抽穗期间温度状况(2013) Table 5 Temperature during heading stage (2013) (℃)

表6 20个杂交组合常温下抽穗期库源性状与耐高温指数的关系(2013) Table 6 Relationship between the sink-source characteristic and high temperature resistance index during heading stage under normal temperature in 20 hybrid combinations (2013)

2.3 人工控制高温对结实率影响及与库源结构和叶片叶绿素含量的关系

上述自然高温条件下研究结果表明, 提高开花当时植株的光合物质供给水平, 有明显减轻高温伤害程度的作用。就同一杂交组合而言, 是否具有同样的效果呢?从试验结果(表7)可见, 随着开花期剪叶面积的增加, 结实率明显下降, 其中高温下的降低幅度更大, 以致开花期剪叶面积与耐高温指数呈极显著负相关; 施氮量越高, 开花期叶绿素含量(SPAD值)和高温下的结实率越高(可能是叶绿素含量高提高了光合速率而增强了其抵御高温胁迫的能力), 但对常温下的结实率没有影响, 以致开花期叶绿素含量与耐高温指数呈显著正相关, 粒叶比与耐高温指数呈极显著负相关(表8)。表明开花期保留较多绿叶面积和较高叶绿素含量对提高耐高温能力作用明显, 与不同组合间粒叶比的表现相符。

表7 不同剪叶程度在高温下的结实率比较(2012, 绵优5240) Table 7 Comparison of seed setting percentage under high temperature and different cutting leaf areas (2012, Mianyou 5240)

表8 不同施氮量在高温下的结实率比较(2013, 冈优169) Table 8 Comparison of seed setting percentage under high temperature and different nitrogen application rates (2013, Gangyou 169)

3 讨论

本研究结果表明, 水稻开花期耐高温组合可分为3类, 一是在高温胁迫下开花具有较强的直接抵御高温能力组合, 属于含耐高温基因的品种^[4]; 二是在每日自然高温发生前完成开花而避开高温直接伤害的品种; 三是源库比较高, 在高温胁迫当时能提供较高的光合物质而缓解高温伤害的品种。以下

针对品种上述耐高温特性就大面积生产如何提高水稻耐高温能力展开讨论。

3.1 关于开花期高温胁迫下的耐高温杂交组合鉴定方法

关于水稻开花期高温胁迫下的生理生化响应的研究较多^{[11, 12, 13, 14, 15, 16, 17, 18, 19, 20, 21]}, 张桂莲等^[11]认为, 水稻耐高温品种的剑叶的光合特性及叶绿素含量、可溶性糖、可溶性蛋白、游离脯氨酸和热稳定蛋白含量较高, 而膜透性和MDA含量较低。任昌福等^[15]、雷东阳等^[16]指出, 高温胁迫下, 光合作用的关键酶活性及叶绿素含量和光合速率均显著下降。汤日圣等^[17]、郭晶心等^[18]、段骅等^[19]的研究结果表明, 开花期高温导致花粉活力、花粉萌发率和结实率下降, 品种间差异明显。这些研究结论对破解品种开花期耐高温的机理和缓解高温伤害相关物化产品的开发均具有重要指导作用。然而, 若要利用这些生理生化指标在高温胁迫下鉴定耐高温品种, 其可操作性极差。结实率是最好的直接指标, 而且方便可靠感观度极强, 较多研究者用耐高温指数=高温下结实率/常温下结实率× 100%作为品种间耐高温性的筛选指标^[5]。但从本试验结果看, 有的组合耐高温指数高, 是由于常温下结实率不高所致。如泰优99高温、常温下结实率分别为72.09%和78.41%, 耐高温指数高达91.94%, 而内5优39高温、常温下结实率分别为77.00%和90.68%, 耐高温指数高达84.91%, 其高温、常温下结实率均明显比泰优99高, 但耐高温指数却比泰优99显著下降(表2)。说明仅用耐高温指数筛选品种有一定缺陷。在生产实践中, 筛选出的耐高温品种用于相应生态区推广, 但高温的发生有一定概率, 正如前述有的耐高温强组合常温下结实率并非很高, 当高温未发生时, 其结实率不高则使高产受到一定限制。因此, 在鉴定耐高温品种时, 还应考虑其在常温下的结实率。本研究中绵香优576、蓉稻优415、香绿优727、绵优5240、川谷优202、川优6203、冈优169、内香优5828、川谷优7329、江优126十个组合无论在常温还是高温条件下抽穗开花获得高产的可能性均较大。

3.2 关于杂交组合开花习性对耐高温特性的影响

谭中和等^[6]在人工气候箱控温条件下的研究结果表明, 在开花后4 h的高温对受精仍有较大影响; 而朱兴明等^[24]在自然高温条件下的研究则认为, 开花期高温伤害的敏感期在开花当时, 这对受精影响最大, 花后1 h则无明显影响, 进一步指出水稻花时随当日气温而变, 气温越高花时越早, 杂交水稻在自然高温下因花时早于日最高温发生前, 受高温伤害较小。从本研究结果看, 35℃以上的高温最早发生在12:30, 而99.48%的植株在12:30以前开花, 高温伤害极小, 但杂交组合间存在显著差异。9:30— 11:00开花比例与耐高温指数呈极显著正相关, 而11:00— 12:30开花比例与耐高温指数呈极显著负相关, 同时受粒叶比影响较大。说明11:00以前开花者受高温伤害较小, 而11:00以后开花者受伤害较大。结实率是鉴定水稻品种开花期耐高温能力最直接的可靠指标, 但需要在高温胁迫条件下完成。为此则需要人工控温或在高温地区通过分期播种手段实施, 而绝大多数单位难以开展此项工作。从本研究结果看, 杂交组合间耐高温能力的差异来自花时的影响, 开花当时光合物质供给能力的不同, 品种间临界温度的不同。因此, 可以把水稻开花习性和库源比作为鉴定避高温伤害能力的重要指标。

3.3 关于提高开花期耐高温能力的栽培管理措施

谭中和等^[6]指出, 川东南高温伏旱区规律性的高温发生在7月下旬至8月上旬, 栽培措施上通过适期早播早栽, 将杂交中籼的齐穗期提早到7月上旬以前避开高温危害, 是解决开花期高温伤害的最佳途径。盛婧等^[9]认为长江流域水稻播种期安排在5月20日之后可避免抽穗期高温危害, 增施穗肥延迟抽穗期的效果极佳。朱兴明等^[24]认为通过栽培方式改善大田植株的通风透光条件, 有利于缓解高温开花期伤害。从近几年高温出现的时间来看有提早的趋势^[3], 但也有推迟的年份, 如2011年川东南的极端高温推迟到了8月中下旬。表明仅通过提早抽穗期避开高温危害并不十分保险, 若加上选用耐高温能力强的组合则更稳妥。

本研究结果表明, 在常温下穗数型杂交组合在开花期自然高温胁迫下有较强的耐高温能力。其原因在于, 有效穗数越高的组合其穗粒数越少, 粒叶比越低, 即单位颖花的光合源占有量越多, 以致抽穗期耐高温指数与粒叶比呈极显著负相关。表明抽穗开花期遇高温情况下, 有充足的光合物质供应有利于减轻高温对颖花受精结实的危害。这与人工控温条件下通过剪叶调节粒叶比及施氮提高叶绿素含量水平的结果一致。因此, 在大面积生产实践中, 通过选用中小穗型的穗数型杂交组合, 增加中期穗肥的施用比例, 提高开花期受高温胁迫当时的叶绿素含量水平及单位颖花的光合源(绿叶面积)占有量, 提高光合产物供给水平, 可显著缓解开花期的高温伤害程度。

4 结论

杂交水稻组合间耐高温能力差异显著, 绵香优576、蓉稻优415、香绿优727、绵优5240、川谷优202、川优6203、冈优169、内香优5828、川谷优7329、江优126十个组合开花期在常温下结实率和耐高温指数均较高, 可作为抽穗期耐热品种大面积推广。有效穗数越少的杂交组合的穗粒数和粒叶比越高, 即单位颖花的光合源占有量越少, 开花期耐高温指数与粒叶比呈显著负相关, 11:00以前开花比例和抽穗期叶绿素含量分别与耐高温指数呈显著正相关。选用穗数型、花时早而集中的杂交组合和提高开花期叶绿素含量水平, 是增强杂交水稻开花期耐高温能力的重要途经。

The authors have declared that no competing interests exist.

作者已声明无竞争性利益关系。The authors have declared that no competing interests exist.

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[16]	雷东阳, 陈立云, 李稳香, 姜琴. 高温对不同杂交稻开花期影响的生理差异. 农业现代化研究, 2005, 26: 397-400 Lei D Y, Chen L Y, Li W X, Jiang Q. Effect of high temperature on physiology difference of flowering among different hybrid rice. Res Agric Modernization, 2005, 26: 397-400 (in Chinese with English abstract) [本文引用:3]
[17]	汤日圣, 郑建初, 张大栋, 金之庆, 陈留根, 黄益洪, 石春林, 葛道阔. 高温对不同水稻品种花粉活力及籽粒结实的影响. 江苏农业学报, 2006, 22: 369-373 Tang R S, Zheng J C, Zhang D D, Jin Z Q, Chen L G, Huang Y H, Shi C L, Ge D K. Effect of high temperature on pollen vitality and seed setting of different rice varieties. Jiangsu J Agric Sci, 2006, 22: 369-373 (in Chinese with English abstract) [本文引用:3] [CJCR: 0.895]
[18]	郭晶心, 曾文智, 周宝津, 卢浩荣, 陈庚平, 唐辉武, 周少川, 刘耀光. 开花期高温胁迫对不同水稻品种花粉萌发和结实的影响. 华南农业大学学报, 2010, 31(2): 50-53 Guo J X, Zeng W Z, Zhou B J, Lu H R, Chen G P, Tang H W, Zhou S C, Liu Y G. Effect of high temperature on pollen germination and seed setting of rice during heading period. J South China Agric Univ, 2010, 31(2): 50-53 (in Chinese with English abstract) [本文引用:3] [CJCR: 0.686]
[19]	段骅, 唐琪, 剧成欣, 刘立军, 杨建昌. 抽穗灌浆早期高温与干旱对不同水稻品种产量和品质的影响. 中国农业科学, 2012, 45: 4561-4573 Duan H, Tang Q, Ju C X, Liu L J, Yang J C. Effect of high temperature and drought on grain yield and quality of different rice varieties during heading and early grain filling periods. Sci Agric Sin, 2012, 45: 4561-4573 (in Chinese with English abstract) [本文引用:3] [CJCR: 1.889]
[20]	李成德. 高温导致水稻出现大量空壳分析. 陕西农业科学, 2003, (5): 45-47 Li C D. Analysis of numerous unfilled grain appeared in rice under high temperature. Shaanxi J Agric Sci, 2003, (5): 45-47 (in Chinese with English abstract) [本文引用:2] [CJCR: 0.3367]
[21]	李天, 刘奇华, 大杉立, 山岸徹, 佐佐木治人. 灌浆结实期高温对水稻籽粒蔗糖及降解酶活性的影响. 中国水稻科学, 2006, 20: 626-630 Li T, Liu Q H, Ryu O, Tohru Y, Haruto S. Effect of high temperature on sucrose content and sucrose-cleaving enzymes activity in rice during grain filling stage. Chin J Rice Sci, 2006, 20: 626-630 (in Chinese with English abstract) [本文引用:2] [CJCR: 1.494]
[22]	徐富贤, 熊洪, 朱永川, 谢戎. 王贵雄. 川东南高温伏旱区杂交中稻超稀栽培对稻米整精米率的影响与组合间库源结构的关系. 植物生态学报, 2005, 29: 829-835 Xu F X, Xiong H, Zhu Y C, Xie R, Wang G X. The effects of cultivation density on the percent of head milled rice and source to sink ratios of mid-season hybrid rice in eastern and southern Sichuan province. Chin J Plant Ecol, 2005, 29: 829-835 (in Chinese with English abstract) [本文引用:3] [CJCR: 1.989]
[23]	徐富贤, 熊洪, 张林, 郭晓艺, 朱永川, 周兴兵, 刘茂. 杂交中稻齐穗后叶片SPAD值衰减对再生力的影响. 中国农业科学, 2009, 42: 3442-3450 Xu F X, Xiong H, Zhang L, Guo X Y, Zhu Y C, Zhou X B, Liu M. Effects of the index decreased of SPAD value for leaf after full heading on ratooning ability. Sci Agric Sin, 2009, 42: 3442-3450 (in Chinese with English abstract) [本文引用:2] [CJCR: 1.889]
[24]	朱兴明, 曾庆曦, 宁清利. 自然高温对杂交稻开花受精的影响. 中国农业科学, 1983, 16(2): 37-44 Zhu X M, Zeng Q X, Ning Q L. Effect of natural high temperature on flowering and fertilization in hybrid rice. Sci Agric Sin, 1983, 16(2): 37-44 (in Chinese with English abstract) [本文引用:3] [CJCR: 1.889]

2007

1.355

0.0

徐小锋, 田汉勤, 万师强. 气候变暧对陆地生态系统碳循环的影响. 植物生态学报, 2007, 31: 175-188

Xu X

, Tian H

, Wan S

Climate warming impacts on carbon cycling in terrestrial ecosystems.

J Plant Ecol, 2007, 31: 175-188 (in Chinese with English abstract)

Climate warming is one key issue of global change and plays an important role on carbon cycling in terrestrial ecosystems. This paper reviews the recent advance in our understanding of global warming and its impacts on terrestrial carbon cycling and underlying mechanisms. We also discuss the state-of-the art in ecosystem modeling and its applications to ecosystem assessment. Climate warming will influence terrestrial carbon cycling in several aspects: 1) net primary productivity (NPP) will decrease in low-latitude region, and increase in mid-and high-latitude zones, totally show an increase on the global scale; 2) soil respiration will increase at the initial stage and then keep relatively stable because of biotic adaptability; 3) plant carbon storage will increase in high-latitude region, and remain stable or even decrease in low-latitude zone, and show a s light increase on the global level; 4) the production and decomposition rate of litterfall will increase; 5) the decomposition rate of soil organic carbon will increase and thus decrease soil carbon stock, meanwhile, soil carbon stock will increase for more carbon input from plant litter. These two processes will trade off in certain degree, resulting in different results for varied ecosystems. On the global scale, soil carbon stock will show a decrease; 6) although the different performances of diverse ecosystems, the global terrestrial ecosystem acts as a weak carbon source. Biophysical, biogeographical and biogeochemical models were developed in the past decades for global change research. In the future research, there are an urgent need to address the interaction among climate warming and other factors including elevated CO 2 , O 3 , drought, fire disturbance. The big challenge we are facing is how to deal with the complexity with multi-factors and multi-scales by using experimental and modeling approaches.

作为全球变化的主要表现之一，气候变暖对全球陆地生态系统碳循环的影响巨大，揭示这一作用对于精确理解碳循环的过程和相关政策的制定具有重要的指导意义。该文综述了此领域近十几年来的主要研究工作，总结了陆地生态系统碳循环对气候变暖响应的主要内部机制及其过程，简述了相关模型的发展及其主要应用，并指出以往研究中存在的主要问题以及未来研究的主要方向。在气候变暖条件下，陆地生态系统碳循环的变化主要体现在以下几个方面：1)低纬度地区生态系统NPP一般表现为降低，而在中高纬度地区通常表现为增加，而在全球尺度上表现为NPP增加；2)土壤呼吸作用增强，但经过一段时间后表现出一定的适应性；3)高纬度地区的生态系统植被碳库表现为增加趋势，低纬度地区生态系统植被碳库变化不大，或略微降低，在全球尺度上表现为植被碳库增加；4) 地表凋落物的产量和分解速率增加；5)土壤有机碳分解加速，进而减少土壤碳储存，同时植被碳库向土壤碳库的流动增加从而增加土壤碳库，这两种作用在不同生态系统的比重不同，在全球尺度上表现为土壤碳库的减少；6)尽管不同生态系统表现各异，总体上全球陆地生态系统表现为一个弱碳源。生物物理模型、生物地理模型和生物地球化学模型陆续被开发出来用于研究工作，并取得了一定的成果，但是研究结果仍然存在很大的不确定性。在未来的数年甚至是数十年间，气候变暖与全球变化的其它表现间的协同影响将是下一步的研究重点，气候变暖和陆地生态系统间的双向反馈作用机制是进行更准确研究的理论基础，生态系统结构和功能对气候变化的适应性是准确理解和预测未来气候情景下陆地生态系统碳循环的前提。

... 引言全球气候变暧已成为一个不争的事实^[1,2], 它将对整个陆地生态系统的碳循环产生重大影响^[1] ...

2011

0.0

... 引言全球气候变暧已成为一个不争的事实^[1,2], 它将对整个陆地生态系统的碳循环产生重大影响^[1] ...

2013

0.0

0.6408

陈升孛, 刘安国, 张亚杰, 吴坤悌. 气候变化背景下湖北省水稻高温热害变化规律研究. 气象与减灾研究, 2013, 36(2): 51-56

Chen S

, Liu A

, Zhang Y

, Wu K

Dynamic variations of heat injury on rice in Hubei Province under climate change.

Meteorol Disaster Reduct Res, 2013, 36(2): 51-56 (in Chinese with English abstract)

Based on the 1951-2010 highest temperature data in Hubei province, the dynamic variations of heat injury are studied by using mathematical statistics method. The results show that the heat injury events on rice occur more frequently in many regions of Hubei province. The occurrence time of maximal heat injury probability presents a significant advance in most area except southwestern Hubei, ahead more than 1 day per 10 years. According to the previous results, the spatial distribution of dynamic variation of heat injury on rice and the degree of risk are zoned by using the ArcGIS software.

利用历年气象资料，运用数理统计方法，分析了湖北省1951-2010年水稻高温热害的动态变化，探讨了气候变化背景下高温热害的演变趋势与规律。结果表明，鄂东部、江汉平原部分地区水稻高温热害发生趋于频繁，且除西南部地区外的湖北省其他地区水稻高温热害最大概率出现的时间均有明显的提前，甚至每10 a提前1 d以上。最后，利用ArcGIS对湖北省的水稻高温热害变化趋势和风险程度进行了区划。

... 水稻是我国南方的主要粮食作物之一, 在全球气候变暖的大环境下, 极端天气频繁发生, 高温已经成为制约水稻产量和品质的主要因素^[3,4] ...

... 从近几年高温出现的时间来看有提早的趋势^[3], 但也有推迟的年份, 如2011年川东南的极端高温推迟到了8月中下旬 ...

2013

0.0

1.1628

李飞, 卓壮, Kapila Siri Udawela U A, 高用明, 石英尧. 水稻高温热害发生机理与耐高温遗传基础研究. 植物遗传资源学报, 2013, 14: 97-103

, Zhuo

, Kapila Siri Udawela U A, Gao Y M, Shi Y Y. Damage mechanism of heat stress and genetic basis of heat tolerance in rice. J Plant Genet Resour, 2013, 14: 97-103 (in Chinese with English abstract)

The extreme climate affairs occur more frequently under the trends of global warming. Heat has been one of the major constrains of rice yield and quality. The most effective way to solve the problem is to develop new rice varieties with heat tolerance. The advance of molecular biotechnology and rice functional genome research provide an effective approach for studying on the genetic bases of rice heat tolerance. The damage mechanism of heat stress and genetic basis of heat tolerance in rice were summarized in this review, e.g. QTL mapping and cloning, transcriptome and proteomics. It may facilitate the deep dissection of genetic mechanism of heat tolerance in rice and the development of new rice variety with heat tolerance.

在全球气候变暖的大环境下，极端天气频繁发生，高温已经成为制约水稻产量和品质的主要因素之一。培育耐高温的水稻品种是防止高温热害最简便有效的途径。现代分子技术的发展和水稻功能基因组研究的深入，为水稻耐高温遗传机制的剖析提供了有效手段。本文综述了近年来水稻高温热害的发生机理和耐高温遗传基础方面的研究进展，包括耐高温QTL定位与克隆、转录组研究、蛋白组研究等。以期为深入剖析水稻耐高温的分子机制和培育耐高温的新品种提供有益参考。

... 水稻是我国南方的主要粮食作物之一, 在全球气候变暖的大环境下, 极端天气频繁发生, 高温已经成为制约水稻产量和品质的主要因素^[3,4] ...

... 因此, 开展水稻品种开花期耐高温机理及其缓解技术研究是近年国内外的热点课题^[4,5,7,8] ...

... 3 讨论本研究结果表明, 水稻开花期耐高温组合可分为3类, 一是在高温胁迫下开花具有较强的直接抵御高温能力组合, 属于含耐高温基因的品种^[4] ...

2007

0.0

1.742

田小海, 松井勤, 李守华, 林俊城. 水稻花期高温胁迫研究进展与展望. 应用生态学报, 2007, 18: 2632-2636

Tian X

, Tsutomu

, Li S

, Lin J

High temperature stress on rice anthesis: research progress and prospects.

Chin J Appl Ecol, 2007, 18: 2632-2636 (in Chinese with English abstract)

水稻花期高温引起的不育危害,具有发生可预测性低、作用局限于特定部位、致害程度严重等特点.本文就水稻花期高温不育问题,在发生时间、致害温度、品种反应差异、作用机制、鉴定方法以及抗性性状遗传等方面作了综述.并结合在日本的相关研究及对中国江汉平原杂交中稻花期高温危害发生的气候条件、品种差异等的初步分析,提出了该领域研究目前存在的关键问题和主要解决途径.

... 特别是抽穗开花期的极端高温造成水稻结实率显著下降而大幅度减产^[5,6] ...

... 因此, 开展水稻品种开花期耐高温机理及其缓解技术研究是近年国内外的热点课题^[4,5,7,8] ...

... 100%作为品种间耐高温性的筛选指标^[5] ...

1985

0.0

1.667

谭中和, 蓝泰源, 任昌福, 方文. 杂交籼稻开花期高温危害及其对策的研究. 作物学报, 1985, 11: 103-108

Tan Z

, Lan T

, Ren C

, Fang

Studies on high temperature injury on hybrid rice at flowering time and the strategy to avoid high temperature damage.

Acta Agron Sin, 1985, 11: 103-108 (in Chinese with English abstract)

Studies were made in phytotron as well as under climatic conditionin southeastern Sichuan in 1982-1983 on the critical high tempe-rature causing sterility to hybrid rice at flowering time,the mechan-ism of high temperature injury and on the index of heat tolerance of twohybrid rice varieties,a strat

1981～1983年在人工气候箱定温和川东南自然高温条件下,研究了杂交籼稻开花期高温致害指标、高温敏感期、高温危害机制,测定了汕优二号和矮优二号开花期耐高温指数,提出了开花期高温危害的对策。研究结果表明:35℃4小时的高温对汕优二号的开花受精具有明显伤害;川东南地区日平均气温≥30℃,日最高气温≥35℃的高温天气对杂交

... 特别是抽穗开花期的极端高温造成水稻结实率显著下降而大幅度减产^[5,6] ...

... 2013年观察到对水稻开花受精具有明显伤害的高温(日最高温度达35℃)^[6]稳定期在7月11日下午 ...

... 2 关于杂交组合开花习性对耐高温特性的影响谭中和等^[6]在人工气候箱控温条件下的研究结果表明, 在开花后4 h的高温对受精仍有较大影响 ...

... 3 关于提高开花期耐高温能力的栽培管理措施谭中和等^[6]指出, 川东南高温伏旱区规律性的高温发生在7月下旬至8月上旬, 栽培措施上通过适期早播早栽, 将杂交中籼的齐穗期提早到7月上旬以前避开高温危害, 是解决开花期高温伤害的最佳途径 ...

2013

0.0

0.9745

王志刚, 王磊, 林海, 庞乾林, 鄂志国, 张玉屏, 朱德峰. 水稻高温热害及耐热性研究进展. 中国稻米, 2013, 19(1): 27-31

Wang Z

, Wang

, Lin

, Pang Q

, E Z G, Zhang Y P, Zhu D F. Research progress on heat damage and heat resistance of high temperature. China Rice, 2013, 19(1): 27-31 (in Chinese)

综述了高温对水稻形态发育、产量、品质、生理生化特性的影响以及水稻耐热性鉴定与高温预测评估的研究进展,并对进一步深入开展水稻耐热性研究提出了一些建议.

... 因此, 开展水稻品种开花期耐高温机理及其缓解技术研究是近年国内外的热点课题^[4,5,7,8] ...

2013

0.0

高伟, 徐延浩, 吴晨阳, 田小海. 水稻生殖生长期的高温危害与表观遗传. 长江大学学报(自然科学版), 2013, 10(11): 1-5

Gao

, Xu Y

, Wu C

, Tian X

High temperature harm at rice reproductive growth stage and epigenetic.

J Yangtze Univ (Nat Sci Edn), 2013, 10(11): 1-5 (in Chinese with English abstract)

从温室效应与水稻(Orgza sativa L．)高温危害、水稻耐热的遗传基础、高温对水稻生理生化的影响、高温对水稻生殖发育的影响、植物有性生殖过程中的表观遗传变化、表观遗传变化与植物逆境应答根等方面综述了水稻生殖生长期的高温危害与表观遗传。

... 因此, 开展水稻品种开花期耐高温机理及其缓解技术研究是近年国内外的热点课题^[4,5,7,8] ...

2006

0.0

0.895

盛婧, 陈留根, 朱普平, 薛新红, 郑建初. 不同水稻品种抽穗期对高温的响应及避热的调控措施. 江苏农业学报, 2006, 22: 325-330

Sheng

, Chen L

, Zhu P

, Sue X

, Zheng J

Responses of rice varieties to high temperature at heading stage and methods for heat escape.

Jiangsu J Agric Sci, 2006, 22: 325-330 (in Chinese with English abstract)

采用盆栽试验研究不同水稻品种(系)在抽穗期对高温的响应,用盆栽和大田种植2种方式研究不同播期处理对水稻的抽穗期避热调控效果,在南京和如皋两地试验田研究不同的增施穗肥处理对水稻抽穗期的影响.结果表明:①在抽穗期高温处理试验中,16个参试水稻品种(系)中有2个品种(系)(Z2001和86优8)表现为耐高温,2个品种(农垦57和武香粳14号)表现为中耐,其余12个品种属高温敏感型;②不同播期处理对水稻抽穗期的调控效果明显,综合历年气象资料,在长江流域水稻生产中,为避免抽穗期高温危害,适播期宜安排在5月20日之后;③通过增加穗肥来延迟水稻抽穗时间的效果甚微.

... 品种选择是抵御高温危害的重要措施^[9,10] ...

... 盛婧等^[9]认为长江流域水稻播种期安排在5月20日之后可避免抽穗期高温危害, 增施穗肥延迟抽穗期的效果极佳 ...

2010

0.0

0.895

石春林, 金之庆, 汤日圣, 郑建初. 水稻颖花结实率对减数分裂期和开花期高温的响应差异. 江苏农业学报, 2010, 26: 1139-1142

Shi C

, Jin Z

, Tang R

, Zheng J

Response difference of seed setting rate of rice florets at the meiosis and anthesis stages to high temperature.

Jiangsu J Agric Sci, 2010, 26: 1139-1142 (in Chinese with English abstract)

减数分裂期和开花期是水稻对高温敏感的2个主要阶段.为明确水稻颖花结实率对减数分裂期和开花期高温的响应差异,本研究以特优559和华粳1号为供试材料,在减数分裂期和开花期,通过设置不同高温水平(31℃、33℃、35℃、37℃、39℃、41℃)和处理时间(1 d、3 d、5 d),分析了高温对水稻颖花结实率的影响.结果表明:减数分裂期随着温度和高温处理时间的增加,颖花日均结实率逐步下降,其规律可用二次曲线拟合,2个品种间响应差异不大;开花期随着温度和高温处理时间的增加,颖花结实率明显下降,其规律可用Logistic曲线拟合,2个品种间响应差异较大.

... 品种选择是抵御高温危害的重要措施^[9,10] ...

2007

0.0

1.889

张桂莲, 陈立云, 张顺堂, 刘国华, 唐文邦, 贺治洲, 王明. 抽穗开花期高温对水稻剑叶理化特性的影响. 中国农业科学, 2007, 40: 1345-1352

Zhang G

, Chen L

, Zhang S

, Liu G

, Tang W

, He Z

, Wang

Effects of high temperature on physiological and biochemical characteristics in flag leaf of rice during heading and flowering period.

Sci Agric Sin, 2007, 40: 1345-1352 (in Chinese with English abstract)

【Objective】To ascertain the physiological reaction and mechanism of adaption to high temperature stress of functional leaves in rice.【Method】For seven day,a study was conducted to determine the effects of high temperature stress(8:00-17:00,37℃,17:00-8:00,30℃)on the chlorophyll content,photosynthetic characteristics,heat stable protein content and membrane permeability of flag leaves in hybrid rice in controlled environment chamber.【Result】The results showed that chlorophyll content and chlorophyll a/b of flag leaves in rice decreased under high temperature stress,increased after high temperature stress,chlorophyll content of high-temperature tolerant cultivar 996 was greater than that of high-temperature sensitive cultivar 4628.The net photosynthetic rate and stomatal conductance of flag leaves declined and substomatal CO2 concentration and transpiration rate of flag leaves increased under high temperature stress,compared with high-temperature sensitive cultivar 4628,high-temperature tolerant cultivar 996 could keep higher photosynthetic characteristics under high temperature stress;Souble sugar and souble protein and free proline content of flag leaves decreased under high temperature stress,but heat stable protein content increased;The increase scope of malonyldialdehyde(MDA)content and membrane permeability in high-temperature sensitive cultivar 4628 was greater under high temperature stress than in high-temperature tolerant cultivar 996.【Conclusion】Under high temperature stress,higher photosynthetic characteristics and chlorophyll content,souble sugar,souble protein,proline,heat stable protein content and lower membrane permeability and MDA content of flag leaves in rice were the physiological basis of high-temperature tolerant cultivar.

【目的】探明水稻功能叶在高温胁迫下的生理反应及适应机理。【方法】在人工气候室对两个耐热性不同的水稻品系进行高温处理（8:00～17:00，37℃，17:00～8:00，30℃）,研究高温胁迫对水稻剑叶叶绿素含量、光合特性、热稳定蛋白和膜透性等理化特性的影响。【结果】高温胁迫下，水稻剑叶叶绿素含量和叶绿素a/b值下降，胁迫后恢复期间稍有回升，耐热品系996叶绿素含量高于热敏感品系4628。高温使剑叶净光合速率和气孔导度下降，细胞间CO2浓度上升，与热敏感品系4628相比，耐热品系996在高温胁迫下能保持较高的光合特性；高温使水稻剑叶中可溶性糖含量、可溶性蛋白质含量和游离脯氨酸含量下降，热稳定蛋白含量上升；热敏感品系4628剑叶中丙二醛含量和膜透性在高温胁迫下增幅大于耐热品系996。【结论】高温胁迫下水稻剑叶中能保持较高的光合特性及叶绿素含量、可溶性糖、可溶性蛋白质、游离脯氨酸和热稳定蛋白含量以及较低的膜透性和MDA含量，是耐高温品种的生理基础。

... 因此, 国内外就水稻在高温胁迫下叶片的光合特性及叶绿素含量、可溶性糖、可溶性蛋白、游离脯氨酸、热稳定蛋白、膜透性、MDA含量和光合作用的关键酶Rubisco、活化酶RuBP、羧化酶活性^{[11,12,13,14,15,16]}, 花粉的活力、萌发率、可育率、受精率、结实率及花器官发育^{[17,18,19,20]}, 籽粒的蔗糖、果糖和葡萄糖含量的动态变化以及蔗糖合酶、液泡型转化酶和细胞壁结合型转化酶活性等^[21]的生理生化响应, 开展了广泛而深入的研究 ...

... 1 关于开花期高温胁迫下的耐高温杂交组合鉴定方法关于水稻开花期高温胁迫下的生理生化响应的研究较多^{[11,12,13,14,15,16,17,18,19,20,21]}, 张桂莲等^[11]认为, 水稻耐高温品种的剑叶的光合特性及叶绿素含量、可溶性糖、可溶性蛋白、游离脯氨酸和热稳定蛋白含量较高, 而膜透性和MDA含量较低 ...

1999

0.0

1.494

黄英金, 罗永锋, 黄兴作, 饶志, 刘宜柏. 水稻灌浆期耐热性的品种间差异及其与剑叶光合特性和内源多胺的关系. 中国水稻科学, 1999, 13: 205-210

Huang Y

, Luo Y

, Huang X

, Rao

, Liu Y

Varietal difference of heat tolerance at grain filling stage and its relationship to photosynthetic characteristics and endogenous polyamine of flag leaf in rice.

Chin J Rice Sci, 1999, 13: 205-210 (in Chinese with English abstract)

The evaluation of heat tolerance at grain filling stage was studied on 65 early season rice varieties from differentgene sources ,and the physiological reaction around high temperature stress at grain filling stage was compared between the varieties with much difference in heat tolerance. The results indicated that the stress index of unfilled grain rate, filled grainweight, head rice rate, chalkiness and protein content could be regarded as the five indexes of heat tolerance for field directevaluation at grain filling stage in rice. There was much difference in heat tolerance among the different varieties according tothe comprehensive evaluation of heat tolerance by subordinate function value. Thus several varieties with high heat tolerancewere screened. The photosynthetic ability of rice can be affected seriously by high temperature stress,but the decrease extentof flag leaf photosynthetic rate of varieties with high heat tolerance is significantly lower. The recovery level is also h igher after removing stress. The changing trend of chlorophyll content, RuBPc activity and endogenous polyamine content is thesame,which is the important physiological characteristics of heat tolerance in rice.

对65个不同来源的早籼稻品种进行了灌浆期耐热性的鉴定研究，并比较了耐热性相差很大的品种灌浆期高温胁迫前后的生理反应。结果表明：秕粒率、实粒重、整精米率、垩白度、蛋白质含量的胁迫指数可作为水稻品种灌浆期耐热性田间直接鉴定的 5个指标，可采用隶属函数值法对各品种的耐热性进行综合评价；不同品种的耐热性存在相当大的差异，筛选出了几个耐热性很强的品种；高温胁迫严重影响水稻的光合能力，但耐热性强的品种剑叶的光合速率高温胁迫后下降幅度明显较小，胁迫解除后恢复的程度也更高，叶绿素含量、RuBPc 活性及内源多胺含量的变化趋势与光合速率相一致，是水稻耐高温逼熟的重要生理特征。

2005

0.0

0.445

朱雪梅, 柯永培, 邵继荣, 林立金, 杨远祥. 高温胁迫对重穗型水稻品种叶片活性氧代谢的影响. 种子, 2005, 24(3): 27-32

Zhu X

, Ke Y

, Shao J

, Lin L

, Yang Y

Effects of high temperature stress on activated oxygen metabolism of the leaves of the heavy panicle type of medium indica hybrid rice.

Seed, 2005, 24(3): 25-32 (in Chinese with English abstract)

利用高温胁迫,比较研究了不同穗重型品种叶片的O-2产生速率、H2O2和MDA含量变化,以及抗坏血酸过氧化物酶(ASP)、过氧化氢酶(CAT)和超氧化歧化酶(SOD)的酶活性变化.结果表明:高温胁迫下,水稻叶片中的O-2产生速率、H2O2和MDA含量随处理时间的延长而加速上升,且上升速率及含量呈现轻穗型品种E优540＞中穗型品种D优68＞重穗型品种II优162、冈优527;而保护酶SOD、CAT和ASP的活性则相反,呈现重穗型品种＞中穗型品种＞轻穗型品种.同时,从高温胁迫下各类型品种叶片中CAT和ASP的活性上升与下降的拐点比较,重穗型品种比中穗型和轻穗型的出现时间推迟1 d.由此认为,重穗型品种的这种抗逆性代谢优势,是该类型在高光强、高温和低CO2浓度下显示其高的净光合速率的生理基础.

2009

0.0

1.667

曹云英, 段骅, 杨立年, 王志琴, 刘立军, 杨建昌. 抽穗和灌浆早期高温对耐热性不同籼稻品种产量的影响及其生理原因. 作物学报, 2009, 35: 512-521

Cao Y

, Duan

, Yang L

, Wang Z

, Liu L

, Yang J

Effect of high temperature during heading and early grain filling on grain yield of indica rice cultivars differing in heat-tolerance and its physiological mechanism.

Acta Agron Sin, 2009, 35: 512-521 (in Chinese with English abstract)

Heading/flowering and early grain filling are the most stress-sensitive periods in rice growth and development. However, little is known how heat-stress during these periods affects the yield formation. This study investigated pollen development, yield components, and some physiological parameters under high temperature treatments during heading and early grain filling. Four indica rice cultivars, Shuanggui 1 and T219 (heat-sensitive), Huanghuazhan and T226 (heat-tolerant), were pot-grown and subjected to high temperature treatments (the mean temperature during the day > 33℃) during heading/flowering (0 – 10 d aft er heading) and early grain filling (11 – 20 d afte r heading), and the natural temperature (the mean temperature during the day The heat-stress treatment significantly increased activities of enzymes involved in antioxidant system in leaves for the hea t-tolerant cultivars, while it much less affected for heat-sensitive cultivars. The high temperature significantly reduced ATPase activity in grains for the heat-sensitive cultivars, and the result was reversed for the heat-tolerant ones. The high temperature increased leaf-temperature and malonyldialdehyde content in leaves ， and reduced root activity and the photosynthetic rate of the flag leaf for both kinds of cultivars, but the increase or reduction was less for heat-tolerant cultivars than for heat-sensitive cultivars. These results suggested that lower leaf-temperature, higher root activity, higher ATPase activity in grains, and higher photosynthetic rate and activities of antioxidant enzymes in leaves would be important physiological traits leading to a higher grain yield for a heat-tolerant rice cultivar.

选用 4 个耐热性不同的籼稻品种，分别于抽穗 ( 始穗后 0 ~ 10 d ) 和灌浆早期 ( 始穗后 11 ~ 20 d) 进行高温 ( 白天温度 >33℃ ) 处理，以同期自然温度 ( 白天温度为对照，研究高温对产量的影响及其生理机制。结果表明，与对照相比，高温处理显著降低热敏感品种的花粉可育率、受精率，而耐热品种与对照无显著差异。高温胁迫明显降低热敏感品种的结实率，导致产量显著下降，且抽穗期高温处理影响大于灌浆早期处理。高温胁迫显著增加耐热品种黄华占叶片的抗氧化酶活性，对热敏感品种影响不大。高温胁迫显著降低热敏感品种籽粒 ATP 酶活性，耐热品种结果则相反。高温处理增加两类品种叶片温度和丙二醛 (MDA) 含量，降低根系活力和叶片光合速率，耐热品种增降的幅度显著小于热敏感品种。在高温胁迫下较低的叶片温度，较强的根系活力和抗氧化保护系统能力及较高的籽粒 ATP 酶活性是耐热性品种保持较高产量的重要生理原因，也是耐热品种的重要生理特征。

1990

0.0

0.45

任昌福, 陈安和, 刘保国. 高温影响杂交水稻开花结实的生理生化基础. 西南农业大学学报, 1990, 12: 440-443

Ren C

, Chen A

, Liu B

The effect of high temperature on the physiology and biochemical foundation of flowering and fertilization of hybrid rice.

J Southwest Agric Univ, 1990, 12: 440-443 (in Chinese with English abstract)

利用人工气候箱研究了高温影响杂交水稻结实率的生理生化基础。结果表明,在29～41℃范围内,叶绿素总量和a、b,总糖、还原糖、蔗糖、糖脂糖、己糖磷酸磷、磷脂磷、高能磷和核酸磷的含量均以32℃为最高,高于32℃时,其含量随温度的升高而下降;无机磷、游离氨基酸总量和游离脯氨酸含量,则随温度的升高而提高。杂交水稻的空秕率以32℃为最低,超过32℃时,则随温度的提高而激剧上升。

... 任昌福等^[15]、雷东阳等^[16]指出, 高温胁迫下, 光合作用的关键酶活性及叶绿素含量和光合速率均显著下降 ...

2005

0.0

雷东阳, 陈立云, 李稳香, 姜琴. 高温对不同杂交稻开花期影响的生理差异. 农业现代化研究, 2005, 26: 397-400

Lei D

, Chen L

, Li W

, Jiang

Effect of high temperature on physiology difference of flowering among different hybrid rice.

Res Agric Modernization, 2005, 26: 397-400 (in Chinese with English abstract)

利用人工气候室研究了高温对不同杂交稻开花期影响的生理差异.结果表明,水稻的耐热性与高温下叶片电导率、游离脯氨酸(pro)、脱落酸(ABA)的增加及叶绿素含量、花粉活力、花药开裂系数的降低有密切的关系.耐热性差的组合叶绿素含量、电导率、花粉活力和花药开裂对高温的响应比耐热性强的迅速,而耐热性强的组合在高温条件下游离脯氨酸和脱落酸含量增加的幅度比耐热性差的大.水稻耐热性与高温条件下叶绿索、游离脯氨酸、脱落酸含量及花粉活力、花药开裂系数呈正相关,与电导率呈负相关.上述6种生理指标可鉴定水稻的耐热性.

... 任昌福等^[15]、雷东阳等^[16]指出, 高温胁迫下, 光合作用的关键酶活性及叶绿素含量和光合速率均显著下降 ...

2006

0.0

0.895

汤日圣, 郑建初, 张大栋, 金之庆, 陈留根, 黄益洪, 石春林, 葛道阔. 高温对不同水稻品种花粉活力及籽粒结实的影响. 江苏农业学报, 2006, 22: 369-373

Tang R

, Zheng J

, Zhang D

, Jin Z

, Chen L

, Huang Y

, Shi C

, Ge D

Effect of high temperature on pollen vitality and seed setting of different rice varieties.

Jiangsu J Agric Sci, 2006, 22: 369-373 (in Chinese with English abstract)

以杂交籼稻汕优63、特优559和常规粳稻武育粳3号、华粳1号为材料,于抽穗期用35℃及其以上高温胁迫处理,研究了高温对花粉活力和籽粒结实的影响.结果表明:35℃胁迫1 d,供试品种的花粉活力、花粉萌发率和结实率均下降;随着胁迫温度的升高和时间的延长,供试品种花粉活力、花粉萌发率和结实率都急剧降低;水稻抽穗期受高温的危害存在品种间差异,相同条件下,杂交籼稻汕优63和常规粳稻武育粳3号受高温的伤害分别明显小于同类型品种特优559和华粳1号;花粉活力降低是高温导致籽粒结实率大幅度降低的主要生理原因.

... 汤日圣等^[17]、郭晶心等^[18]、段骅等^[19]的研究结果表明, 开花期高温导致花粉活力、花粉萌发率和结实率下降, 品种间差异明显 ...

2010

0.0

0.686

郭晶心, 曾文智, 周宝津, 卢浩荣, 陈庚平, 唐辉武, 周少川, 刘耀光. 开花期高温胁迫对不同水稻品种花粉萌发和结实的影响. 华南农业大学学报, 2010, 31(2): 50-53

Guo J

, Zeng W

, Zhou B

, Lu H

, Chen G

, Tang H

, Zhou S

, Liu Y

Effect of high temperature on pollen germination and seed setting of rice during heading period.

J South China Agric Univ, 2010, 31(2): 50-53 (in Chinese with English abstract)

Effect of high temperature on pollen germination and seed setting of five rice cultivars during heading period was investigated.The results showed that the pollen germination rate of Huanghuazhan under high temperature for 4 d was above 70%,and its seed setting rate under high temperature 7 d was about 50%,while the other cultivars had lower pollen germination rates and seed setting rates.Therefore,Huanghuazhan is a heat-tolerant cultivar.

以5个籼稻品种为材料,通过抽穗期高温处理和离体花粉萌发等试验,研究了开花期高温对水稻花粉萌发和结实的影响.结果表明,不同品种水稻对高温处理的敏感度明显不同,其中品种黄华占在高温处理4 d后的花粉萌发率达到70%以上,在高温处理7 d后小穗结实率仍保持50%左右,其他4个品种的花粉萌发率和小穗结实率都明显低于黄华占,由此认为,黄华占是一个耐热性较强的品种.

... 汤日圣等^[17]、郭晶心等^[18]、段骅等^[19]的研究结果表明, 开花期高温导致花粉活力、花粉萌发率和结实率下降, 品种间差异明显 ...

2012

0.0

1.889

段骅, 唐琪, 剧成欣, 刘立军, 杨建昌. 抽穗灌浆早期高温与干旱对不同水稻品种产量和品质的影响. 中国农业科学, 2012, 45: 4561-4573

Duan

, Tang

, Ju C

, Liu L

, Yang J

Effect of high temperature and drought on grain yield and quality of different rice varieties during heading and early grain filling periods.

Sci Agric Sin, 2012, 45: 4561-4573 (in Chinese with English abstract)

【Objective】Heading/flowering and early grain filling are the most stress-sensitive periods in rice growth and development. However, little is known about how high temperature and water stress during these periods affect rice yield and grain quality. This study investigated pollen development, yield components, grain quality and some physiological parameters under high temperature and water stress during heading and early grain filling periods.【Method】Two conventional indica rice cultivars, Shuanggui 1 (heat-sensitive) and Huanghuazhan (heat-tolerant), and two super rice cultivars, Liangyoupeijiu (indica-inclined two-line hybrid) and Yangjing 4038 (japonica), were grown in pots and subjected to high temperature treatments (HT, maximum temperature during the day at 37.5℃), soil drying treatments (SD, soil water potential at -30±10 kPa) and high temperature plus soil drying treatments (HT+SD), and the natural temperature plus well-watered treatment was taken as the control (CK).【Result】The results showed that high temperature, soil drying or high temperature plus soil drying treatments significantly reduced pollen fertility rate, fertilization rate, seed-setting rate, grain yield, milled rice, head rice and break down viscosity, increased chalky grains, chalkiness degree and setback viscosity in each cultivars. The effect was more obvious on heat-sensitive than on heat-tolerant cultivars, and more evident on Liangyoupeijiu and Yangjing 4038 than on Shuanggui 1 and Huanghuazhan under both high temperature and high temperature plus soil drying treatments. The result was discrepant under soil drying treatments, the extent of the decrease or increase was more on heat-sensitive than on heat-tolerant cultivars, but more evident on Shuanggui 1 and Huanghuazhan than on Liangyoupeijiu and Yangjing 4038. The effect of high temperature treatments, soil drying treatments or high temperature plus soil drying treatments on yield and grain quality was consistent with that on root activity, net photosynthetic rate of the flag leaf and activities of antioxidative enzymes.【Conclusion】The results suggest that the reduction of grain yield and grain quality under high temperature or soil drying treatments during heading and early grain filling varies largely with rice cultivars. Heat-tolerance is better for Shuanggui 1 and Huanghuazhan than for Liangyoupeijiu and Yangjing 4038, and drought-tolerance is stronger for Liangyoupeijiu and Yangjing 4038 than for Shuanggui 1 and Huanghuazhan. Stronger root activity and antioxidative defense system and greater net photosynthetic rate of the flag leaf would be an important physiological mechanism in maintaining a higher grain yield for a rice variety under high temperature or drought stress.

【目的】探讨在高温和干旱胁迫下不同水稻品种产量和品质形成的特点。【方法】常规中熟籼稻品种黄华占（耐热品种）和双桂1号（热敏感品种）与超级稻品种两优培九（两系杂交籼稻）和扬粳4038（粳稻）种植于盆钵，于抽穗和灌浆早期（始穗后0—10 d）设置高温（日最高温度为37.5℃）、干旱（土壤水势保持在-30±10 kPa）、高温+干旱3种处理，以同期自然温度且正常供水为对照。【结果】与对照相比，高温、干旱和高温+干旱3种处理均显著降低了各品种的花粉可育率、受精率、结实率和产量；降低了精米率、整精米率和崩解值，增加了垩白米率、垩白度和消减值。高温及高温+干旱处理对产量和稻米品质的影响，热敏感品种双桂1号大于耐热品种黄华占，两优培九和扬粳4038大于双桂1号和黄华占。干旱对产量和稻米品质的影响，热敏感品种大于耐热品种；双桂1号和黄华占大于两优培九和扬粳4038。高温、干旱或高温+干旱胁迫对产量和稻米品质的影响与其对根系氧化力、叶片光合速率和抗氧化保护酶活性的影响结果趋势一致。【结论】抽穗与灌浆早期高温或干旱胁迫对水稻产量和品质的影响在品种间存在很大差异，双桂1号和黄华占的耐热性强于两优培九和扬粳4038，两优培九和扬粳4038的耐旱性则强于双桂1号和黄华占。在高温或干旱胁迫下根系氧化力、叶片光合速率和抗氧化保护酶活性高是一些水稻品种保持较高产量的重要生理原因。

... 汤日圣等^[17]、郭晶心等^[18]、段骅等^[19]的研究结果表明, 开花期高温导致花粉活力、花粉萌发率和结实率下降, 品种间差异明显 ...

2003

0.0

0.3367

李成德. 高温导致水稻出现大量空壳分析. 陕西农业科学, 2003, (5): 45-47

Li C

Analysis of numerous unfilled grain appeared in rice under high temperature.

Shaanxi J Agric Sci, 2003, (5): 45-47 (in Chinese with English abstract)

高温导致水稻出现大量空壳植株的基本特征是:只有空壳和实粒之分,几乎没有秕粒,茎秆上出现高位分枝,有包颈现象,植株衰老延缓,除高温这一主导因素外,品种、地理位置和地形对空壳的产生有一定影响.选用耐高温品种,合理安排育秧、插秧时间,抽穗期避开高温,是主要防止措施.实施健身栽培,有助于降低空壳率.当高温出现时,通过适当的排灌水技术和喷施微肥,可以减少空壳损失.

2006

0.0

1.494

李天, 刘奇华, 大杉立, 山岸徹, 佐佐木治人. 灌浆结实期高温对水稻籽粒蔗糖及降解酶活性的影响. 中国水稻科学, 2006, 20: 626-630

, Liu Q

, Ryu

, Tohru

, Haruto

Effect of high temperature on sucrose content and sucrose-cleaving enzymes activity in rice during grain filling stage.

Chin J Rice Sci, 2006, 20: 626-630 (in Chinese with English abstract)

The dynamic changes of sucrose,fructose,glucose contents and differences in sucrose synthase(SS),vacuolar invertase(VCI),and cell wall bound invertase(CWI) activities in rice grain after flowering were studied under natural and high temperatures by using two japonica varieties Koshihikari and Sasanishiki as materials.In rice grains,sucrose synthase activity was higher than invertase activity,and positively significantly correlated with starch accumulation rate,indicating that sucrose synthase played an important role in sucrose degradation and starch synthesis.Sucrose content in rice grains(increased) significantly,whereas fructose and glucose contents did not increase correspondingly under high temperature,suggesting that high temperature treatment enhanced sucrose accumulation,but diminished sucrose degradation in rice grains.Sucrose synthase,vacuolar invertase,and cell wall bound invertase activities in rice grains at high temperature decreased definitely compared with controls,showing that the deceleration of sucrose-degradation rate was related to the decrease of(sucrose) synthase and invertase activities.

选用水稻品种越光和笹锦为材料，在开花后设置自然温度和高温两种处理，研究了不同温度处理下籽粒蔗糖、果糖和葡萄糖含量的动态变化以及蔗糖合酶、液泡型转化酶和细胞壁结合型转化酶活性的差异。高温处理下，蔗糖合酶活性值大于转化酶活性值，且与淀粉积累速率呈显著正相关，表明蔗糖合酶在蔗糖的分解和淀粉的合成过程中起着重要作用；高温下两品种籽粒的蔗糖含量明显增加，但分解的葡萄糖和果糖含量并未相应增加，表明高温有利于籽粒中蔗糖的积累，而不利于籽粒中蔗糖的分解。高温下蔗糖合酶、液泡型转化酶和细胞壁结合型转化酶活性明显下降，表明蔗糖分解速率的下降与蔗糖合酶和转化酶活性下降有关。

2005

0.0

1.989

徐富贤, 熊洪, 朱永川, 谢戎. 王贵雄. 川东南高温伏旱区杂交中稻超稀栽培对稻米整精米率的影响与组合间库源结构的关系. 植物生态学报, 2005, 29: 829-835

Xu F

, Xiong

, Zhu Y

, Xie

, Wang G

The effects of cultivation density on the percent of head milled rice and source to sink ratios of mid-season hybrid rice in eastern and southern Sichuan province.

Chin J Plant Ecol, 2005, 29: 829-835 (in Chinese with English abstract)

Many studies on grain quality as affected by rice cultivation densityhave been conducted on one or two rice varieties, but there are few studies on varieties of hybrid midseason_rice. The hybrid midseason_rice grown in southern and eastern Sichuan Province has very low grain quality, especially the percentage of head milled rice, because of high temperatures and summer droughts. This paper explores the effects of low density cultivation on the percentage of head milled rice and source to sink ratios of varieties of hybrid midseason_rice, andit provides a theoretical and practical basis for improved cultivation. The experiment was conducted with eighteen mid_season rice hybrids grown at two densities, a low density cultivation and a traditional cultivation system, ina randomized block design with 4 replications using two hybrids (a big_panicletype hybrid `Gangyou 22' and a small_panicle type hybrid `II you 7') with 3 different treatments of grain or leaf cut. By using variance, correlation and regre ssion analysis, the results indicated that the percentage of head milled rice in creased in hybrids with small_medium panicles under low density cultivation butnot for the hybrids with big_panicles. This was because of the significant decrease in the leaf to grain ratio and grain filling rate of hybrids for small_middle panicle under low density cultivation, as compared to the traditional cultivation system, resulting in an increase in the grain unit weight and percentage ofhead milled rice. The leaf to grain ratio and the grain filling rate of big panicle hybrids were low in traditional cultivation, and decreased significantly under low density cultivation, causing a decrease in the grain unit weight and percentage of head milled rice. These results were produced because of an imbalancein the leaf to grain ratios. Hybrids with less than 177 spikelets per panicle un der cultivation using traditional densities can adapt to the low density cultivation in the eastern and southern regions of Sichuan Province.

以18个杂交中稻组合材料，研究了超稀栽培与传统栽培间稻米整精米率的变异与组合间库源结构的关系。结果表明：中小穗型组合在超稀栽培条件下的整精米率会明显改善,而大穗型组合的整精米率反而降低。其原因在于中小穗型组合在传统栽培条件下的叶粒比和灌浆速率较高，改为超稀栽培后，叶粒比和灌浆速率明显降低，相对提高了籽粒容重，近而提高了整精米率。大穗型组合在传统栽培条件下的叶粒比和灌浆速率较低，改为超稀栽培后, 叶粒比和灌浆速率进一步明显降低，此时因库源比例失调，源明显不足以致籽粒充实度差，从而相对降低了籽粒容重和整精米率。超稀栽培应选用传统栽培条件下着粒数不超过177粒的组合为宜。

... 本田苗期采用常规方法考查最高苗数, 齐穗期采用干物重法考查3月5日播种的各杂交组合的粒叶比^[22], 成熟期各播期所有组合的每次重复均取样5穴, 考查穗部性状, 抽穗期的耐高温指数=高温下结实率/常温下结实率#cod#x000D7 ...

... 齐穗期将3月5日播期(常温下抽穗)中与3月25日(高温下抽穗)挂牌标记相同的20个组合, 各取样5穴, 3次重复, 考查粒叶比^[22]、顶部3叶叶绿素含量(SPAD值)^[23]和单茎干物重 ...

... 于成熟期分别收获, 考查各处理结实率和耐高温指数, 其中2013年试验在移至人工气候室高温处理前测顶部3片叶的叶绿素含量(SPAD值)^[23]和粒叶比^[22] ...

2009

0.0

1.889

徐富贤, 熊洪, 张林, 郭晓艺, 朱永川, 周兴兵, 刘茂. 杂交中稻齐穗后叶片SPAD值衰减对再生力的影响. 中国农业科学, 2009, 42: 3442-3450

Xu F

, Xiong

, Zhang

, Guo X

, Zhu Y

, Zhou X

, Liu

Effects of the index decreased of SPAD value for leaf after full heading on ratooning ability.

Sci Agric Sin, 2009, 42: 3442-3450 (in Chinese with English abstract)

[目的] 探明影响再生力的主作关键植株性状,为杂交水稻及其再生稻的高产育种与栽培提供参考.[方法] 以18个杂交中稻品种为材料,在大田高产栽培条件下,通过相关、回归与通径分析,研究了杂交中稻植株性状与再生力关系及主作和再生总体高产组合的穗粒结构.[结果] 活芽率、发苗力、再生稻有效穗和产量4个性状是代表品种再生力的关键因子,增加有效穗是进一步提高再生稻产量的重要途径;穗粒数、叶颖花比、叶粒数比、叶粒重比、LAI衰减指数和SPAD值衰减指数分别与活芽率,发苗力、再生稻有效穗和产量呈显著或极显著相关,其中仅有头季稻齐穗期至成熟期叶片SPAD值衰减指数对活芽率、发苗力、再生稻有效穗和产量的偏相关均达显著以上水平;同一个组合同时实现头季稻和再生稻均分别达到最高产量的可能性小,两季总产同时满足理论产量达1I.5 t·hm-2和产量潜力达14 t·hm-2的高产组合的主作群体主要特征最佳取值范围:穗粒数为160～190粒,叶粒重比0.0737～0.0827 cm2·mg-1.SPAD值衰减指数0.4029～0.5409、有效穗232.12万～249.40万/hm2、结实率81.54%～85.74%、千粒重28.58～30.07 g、单穗重4.13～4.43 g.[结论] 再生稻进一步高产的主攻目标是增加有效穗,头季稻齐穗到成熟叶片SPAD值衰减指数可作为鉴定再生力的新指标,中等偏大穗品种是中稻-再生稻总体高产的重要特征.

1983

0.0

1.889

朱兴明, 曾庆曦, 宁清利. 自然高温对杂交稻开花受精的影响. 中国农业科学, 1983, 16(2): 37-44

Zhu X

, Zeng Q

, Ning Q

Effect of natural high temperature on flowering and fertilization in hybrid rice.

Sci Agric Sin, 1983, 16(2): 37-44 (in Chinese with English abstract)

正开花期过高温度对水稻开花受精的影响,国内外早有明确的研究结果。但这些试验是在人工恒定温度条件下进行的,故试验结果不能完全反映自然变温条件下高温对水稻开花受精的影响。我国杂交稻推广以来,一些地区鉴于杂交稻抽穗结实期常遇高温天气,又先后开展了高温对杂交稻开花结实影响的试验,但在对自然高温影响的大小和影响实质的评价上,尚存在不同看法。有鉴于此,我们针对四川东南部高温伏旱区的特定高温天气条件,进行了自然高温对杂交稻开花受精影响的观察,为杂交稻在本区的合理布局和栽培提供依据。

... 分析其原因, 开花期高温伤害的敏感期在开花当时, 对受精影响最大, 花后1 h则无明显影响^[24], 而本试验35 ℃以上的日最高温出现在12:30以后, 9:30#cod#x02014 ...

... 而朱兴明等^[24]在自然高温条件下的研究则认为, 开花期高温伤害的敏感期在开花当时, 这对受精影响最大, 花后1 h则无明显影响, 进一步指出水稻花时随当日气温而变, 气温越高花时越早, 杂交水稻在自然高温下因花时早于日最高温发生前, 受高温伤害较小 ...

... 朱兴明等^[24]认为通过栽培方式改善大田植株的通风透光条件, 有利于缓解高温开花期伤害 ...