Enclosed stigma contributes to higher spikelet fertility for rice(Oryza sativa L.) subjected to heat stress

With global warming, rice plants may be subjected to heat stress more regularly during the heatsensitive flowering stage, causing spikelet sterility and grain yield loss.Stigma exsertion is considered to increase pollen reception and promote female reproductive success.The aim of this study was to investigate the role of stigma exsertion on spikelet fertility at high temperatures.Five rice cultivars(Liangyoupeijiu, Shanyou 63, Huanghuazhan, Nagina 22, and IR64) with differing degrees of stigma exsertion were cultivated and exposed to high temperature at anthesis.Heat-tolerant cultivars did not always show a high percentage of spikelets with exserted stigmas, and vice versa.Irrespective of the presence of more pollen grains on exserted stigmas, spikelets with exserted stigmas did not show greater spikelet fertility than spikelets with fewer exserted stigmas or hidden stigmas under heat stress.GA3 application augmented the percentage of spikelets with exserted stigmas;however, it did not increase spikelet fertility under heat stress.Spikelet fertility of whole panicles was negatively correlated with the percentage of spikelets with exserted stigmas, but positively with that with hidden stigmas.Viability of the hidden stigmas was less reduced than that of exserted stigmas under heat stress, suggesting that hidden stigmas have an advantage in maintaining viability.Heat stress delayed anther dehiscence and reduced the viabilities of both exserted stigmas and pollens, thereby causing low spikelet fertility.Together, these results suggest that high spikelet fertility does not depend on stigma exsertion and that enclosed stigma generally contributes to higher spikelet fertility and heat tolerance under high-temperature conditions during flowering in rice.     

The Early-Morning Flowering Trait of Rice Reduces Spikelet Sterility under Windy and Elevated Temperature Conditions at Anthesis

Abstract

We previously demonstrated that heat-induced spikelet sterility at anthesis could be mitigated by using an early-morning flowering (EMF) line of Oryza sativa L. cv. Koshihikari crossed with wild rice, O. officinalis. Another microclimate factor, high wind velocity, is known to increase the sterility induced by heat. In this study, we evaluated whether EMF rice could mitigate sterility under the combined stresses of heat and wind. Rice plants were exposed to three levels of wind velocity (1.1, 2.2, 3.4 m s^?1) from early-morning until 1500 in a glasshouse, where air temperature reached 30°C at 0800, 34°C at 1000 and 38°C around noon. Under these conditions, sterility steadily increased in Koshihikari, ranging from 28.4 to 86.9% as wind velocity increased. However, in the EMF line, low levels of sterility were observed since most spikelets flowered before 1000 when air temperature reached 35°C, the critical value for causing sterility. These results indicated that the increase in heat-induced spikelet sterility by wind can be potentially mitigated by using EMF rice.     

超级杂交稻国稻6号对开花结实期高温热害的反应

以超级杂交稻国稻6号（内2优6号）为材料,于2005-2006年在杭州分期播种,通过不同齐穗期条件下的产量及产量构成因子变异研究,分析开花结实期自然高温对灌浆结实的影响,同时在温室条件下设置可控极值高温（40～42℃）测定热害指数。国稻6号对开花结实期自然高温或设计极值高温的反应明显较对照协优46钝感,两者在小穗育性和热害指数上的差异达显著水平（P<0.05）。试验结果还表明国稻6号随花期日平均温度升高小穗不育率增加,尤其与日最高温度的关系更为密切,两者的相关系数分别为0.8604和0.9850（P<0.05）,表明日最高温度对小穗育性伤害更烈。而灌浆结实期日最高温度对国稻6号结实率的影响不如日平均温度,相关系数分别为0.7352和0.9317（P<0.05）。     

Research Progress on Heat Stress of Rice at Flowering Stage

Global warming has caused frequent occurrence of heat stress at the flowering stage of single-season rice in the Yangtze River region of China, which results in declines of spikelet fertility and yield in rice. Rice flowering stage is the most sensitive period to high temperatures, and therefore, the key for heat stress happening is the flowering stage coinciding with high temperature, which causes spikelet fertility decreasing in heat-sensitive varieties, and is the major factor for heat injury differences among various rice planting regions. With the development of rice breeding, temperature indexes for heat stress has been converted from daily maximum temperature of 35 °C to 38 °C with the stress duration of more than 3 d. During the flowering stage, anther dehiscence inhibition and low pollen shedding onto the stigma are two main reasons for spikelet fertility reduction under high temperatures. At panicle initiation stage, high temperatures aggravate spikelet degeneration, and destroy floral organ development. Various types of rice varieties coexist in production, and indica-japonica hybrid rice demonstrates the highest heat resistance in general, followed by indica and japonica rice varieties. In production, avoiding high temperature is the main strategy of preventing heat stress, and planting suitable cultivars and adjustment of sowing date are the most effective measures. Irrigation is an effective real-time cultivation measure to decline the canopy temperature during the rice flowering stage. We suggested that further study should be focused on exploring heat injury differences among different rice variety types, and innovating rice-planting methods according to planting system changes in rice planting regions with extreme heat stress. Meanwhile, high temperature monitor and warning systems should be improved to achieve optimal heat stress management efficiencies.     

Effects of salt and low light intensity during the vegetative stage on susceptibility of rice to male sterility induced by chilling stress during the reproductive stage

We tested whether exposing rice plants to abiotic stress (salt or shade) during vegetative growth affects the chilling tolerance of reproductive organs, which is one of the most important traits for rice growing in a cool climate; we used two rice cultivars with different tolerance in two growing seasons. We divided the vegetative growth into three phases to clarify the most sensitive period: 7–22 days after transplanting (DAT), 23–38 DAT and 39–54 DAT. Chilling tolerance of the pre-stressed plants was based on the male sterility induced by low temperatures. Shade and salt stress during all three vegetative growth phases significantly reduced stomatal conductance. Shade decreased the specific leaf weight and the leaf sugar and starch contents, but salt had no significant effect, despite causing leaf damage. Low temperatures during the reproductive stage induced spikelet sterility in all plants, but the magnitude was greater in the salt- and shade-stressed plants of both cultivars, especially those stressed late during vegetative growth. The increased spikelet sterility caused by chilling was closely related to the reduction of the total spikelet number per panicle. This is the first study to show that salt and low light stress during vegetative growth increased the susceptibility of rice plants to chilling damage during panicle development.     

Cropping calendar options for rice-wheat production systems at high-altitudes

The onset of rains during dry to wet transition fallow periods in rice-wheat production systems in Nepal cause substantial losses of soil nitrogen if the system is improperly managed. To make use of available nutrients and water, this transition period can either be shortened by early rice planting, or be extended by late planting, allowing a third crop to be grown. Shifting planting dates would require rice genotypes adapted to the different environments. Crop duration is influenced by both vegetative and reproductive development, which in turn is influenced by the photo-thermal environment and genotypic responses to it. An experiment was conducted to derive genotypic photo-thermal constants from phenological observations on diverse rice cultivars, which were then applied to the concept of the phenological model RIDEV to design cropping calendar options. Environmental effects on variation of crop duration were determined by planting at different dates. The risk of yield losses to sterility caused by low temperatures was estimated by simulation. Thirty-one different genotypes of rice were planted at 8 dates in 15-day intervals starting 27 April 2004 at the experimental field of the Regional Agriculture Research Station, Lumle, Nepal. The shortest duration to flowering was observed for planting dates in late May and early June. Simulation of flowering dates with RIDEV yielded correct results only for the early planting dates. For later planting dates simulated flowering dates showed an increasing deviation from the observed. In most cultivars, minimum air temperature below 18 °C during booting to heading stages caused near-total spikelet sterility and a specific delay in flowering. However, the chilling tolerant cultivars Chomrong and Machhapuchre-3 cultivated at high altitude showed less than 30% spikelet sterility even at 15 °C. Simulating crop durations with the derived thermal constants allowed evaluating the different calendar options for high altitude systems.     

Screening for Spikelet Fertility and Validation of Heat Tolerance in a Large Rice Mutant Population

A total of 10 000 M4 individuals in Jao Hom Nil (JHN) mutant population was treated with high temperature (40 ℃ to 45 ℃) during the day time (6 h) from the booting to the harvesting stages, and ambient temperature (33 ℃ to 35℃) was used as the control. The results of screening and yield trials found that the mutant line M9962 had a high spikelet fertility of 78% under heat stress. In addition, the other mutant lines, including M3181 and M7988, had a spikelet fertility of approximately 70%. However, the JHN wild type, Sin Lek, RD15 and RD33 had very low spikelet fertility of 34%, 14%, 9% and 4%, respectively. The lower spikelet fertility at an elevated temperature resulted in a dramatic decrease of filled grain and contributed to a loss in 100-grain weight. M9962 is a potential genetic stock for use in a heat tolerance breeding programme. In addition, spikelet fertility at high temperature was representative of heat tolerance and can be used as a screening trait for heat tolerance during the reproductive phase on a large scale.     

Species,ecotype and cultivar differences in spikelet fertility and harvest index of rice in response to high temperature stress

Spikelet fertility (seed-set) is an important component of yield that is sensitive to high temperature. The objectives of this research were (a) to quantify the effects of high temperature on spikelet fertility and harvest index of rice; (b) to determine if there were species, ecotype, and/or cultivar differences in response to high temperature; and (c) to understand the reasons for lower and/or differential spikelet fertility and harvest index of rice cultivars at high temperatures. Fourteen rice cultivars of different species (Oryza sativa and Oryza glaberrima), ecotypes (indica and japonica) and origin (temperate and tropical) were exposed to ambient and high temperature (ambient + 5 °C) at Gainesville, Florida. High temperature significantly decreased spikelet fertility across all cultivars, but effects varied among cultivars. Based on decreases in spikelet fertility at high temperature, cultivar N-22 was most tolerant, while cultivars L-204, M-202, Labelle, Italica Livorna, WAB-12, CG-14 and CG-17 were highly susceptible and cultivars M-103, S-102, Koshihikari, IR-8 and IR-72 were moderately susceptible to high temperature. There were no clear species or ecotype differences, as some cultivars in each species or within ecotypes of tropical and temperature origin were equally susceptible to high temperature (for example M-202 temperate japonica, Labelle tropical japonica, CG-14 O. glaberrima, and WAB-12 interspecific). Decreased spikelet fertility and cultivar difference at high temperature were due mainly to decreased pollen production and pollen reception (pollen numbers on stigma). Lower spikelet fertility at elevated temperature resulted in fewer filled grains, lower grain weight per panicle, and decreased harvest index. There is a potential for genetic improvement for heat tolerance, thus it is important to screen and identify heat-tolerant cultivars. Spikelet fertility at high temperature can be used as a screening tool for heat tolerance during the reproductive phase.     

水稻高温败育模拟模型

在温度控制试验和文献查阅的基础上,分析了水稻开花期开花概率密度分布、日开花分布规律以及结实率的变化规律,提取了高温对水稻结实的影响因子。开花概率密度随开花历期呈Possion分布,日开花结实率可用二次方程描述,日开花量呈正态分布,高温对开花结实率的影响可用S型曲线描述。进一步结合气温日变化规律,并综合考虑高温对已开和未开颖花的影响,构建了水稻高温败育模拟模型。利用独立的试验资料对模型进行了检验。结果表明,当温度低于40℃时,模拟值与实测值有较好的一致性;当温度高于40℃时,因高温对开放颖花的结实率仍有影响,模拟值与实测值有一定误差。总体上,模拟值与实测值的均方根差（RMSE）为0.12。由于我国大部分地区水稻开花期的日极端最高气温较少超过40℃,因此所建立的模型可较好地模拟高温对水稻结实率的影响。     

非洲栽培稻垩白粒率耐热性QTL的定位

【目的】本研究旨在定位一个稻米垩白粒率高温耐性QTL,为外观品质育种及解析垩白粒率高温耐性的遗传机制提供依据。【方法】以非洲栽培稻耐热品种IRGC102309(Oryza glaberrima Steud.)和籼稻品种R9311(O. sativa L. subsp. indica Kato.)为亲本构建的栽培稻种间染色体片段导入系CSIL05-23为材料构建次级分离群体,结合人工气候室模拟灌浆期高温胁迫处理,采用垩白粒率高温钝感值为评价指标,对非洲栽培稻垩白粒率高温耐性 QTL 进行检测。【结果】 在BC₆F₂分离群体,利用单标记分析,发现第5染色体上的SSR标记RM1200与垩白粒率耐热性状极显著正相关（P=0.0005）。进一步利用BC₆F₃和BC₆F₄分离群体,采用QTL Cartographer 2.5软件和复合区间作图法在水稻第5染色体上的SSR标记RM1200-RM5796区间重复检测到一个灌浆期垩白粒率耐热性QTL, 命名为qHTCGR5,分别解释11.4%和17.5%表型变异。根据BC₆F₄分离群体的纯合重组体表型分组,利用置换作图方法将目标QTL同样定位在SSR标记RM1200-RM5796之间,遗传图距为1.3 cM,物理图距约为333.4 kb。【结论】 控制垩白粒率耐热性的qHTCGR5是一个能够用于稻米外观品质育种的新QTL。     

Characterization of physicochemical qualities and starch structures of two indica rice varieties tolerant to high temperature during grain filling

An extreme high temperature during grain filling is an important environmental factor that reduce the yield and quality of rice. In this study, we compared the grain appearance, composition and starch structure of four rice varieties response to high temperature during grain filling. The results obviously revealed that two indica AUS varieties Halwa and Jamir showed more tolerance to high temperature during grain filling than either the japonica Nipponbare or another AUS rice DJ24. The data showed that Halwa and Jamir presented less chalkiness, less grain weight loss, less change of endosperm components as well as starch structure than Nipponbare and DJ24 under high temperature treatment. Hence, the results implied that Halwa and Jamir might be useful targets to further identify genetic mechanism response to high temperature, at least during grain filling and endosperm development. Moreover, these AUS varieties have potential application value in further breeding of rice with good quality and heat tolerance.     

Research Progress and Breeding Prospects of Grain Size Associated Genes in Rice

The breeding objectives of rice include high yield and good quality. Grain size is the three-dimensional structure of seeds, which is one of the key traits determining grain yield and quality in rice. To date, more than 60 genes associated with grain size have been cloned, but the application of grain size genes in rice breeding has not been summarized. This review covers the current advances which can deepen our understanding, such as the key genes for grain size, genetic mechanism behind this trait, breeding application of cloned genes, functional markers for molecular breeding and prospects for future study, which will provide information of molecular breeding for rice breeders.     

Mapping QTL for Heat-Tolerance at Grain Filling Stage in Rice

A mapping population of 98 lines (backcross inbred lines, BILs) derived from a backcross of Nipponbare/Kasalath// Nipponbare was planted at two experimental sites, Nanjing and Nanchang, and treated with high and optimal temperature during grain filling, respectively. The grain weight heat susceptibility index [GWHSI= (grain weight at optimum temperature-grain weight at high temperature) / grain weight at optimum temperature ×100] was employed to evaluate the tolerance of rice to heat stress. A genetic linkage map with 245 RFLP markers and a mixed linear-model approach was used to detect quantitative trait loci (QTLs) and their main effects, epistatic interactions and QTL×environment interactions (Q×E). The threshold of LOD score=2.0 was used to detect the significance of association between marker and trait. A total of 3 QTLs controlling heat tolerance during grain filling were detected, on chromosomes 1, 4 and 7, with LOD scores of 8.16, 11.08 and 12.86, respectively, and they explained the phenotypic variance of 8.94, 17.25 and 13.50 %, correspondingly. The QTL located in the C1100-R1783 region of chromosome 4 showed no QTL×environment interaction and epistatic effect, suggesting that it could be stably expressed in different environments and genetic backgrounds, and thus it would be valuable in rice breeding for heat tolerance improvement. This QTL allele, derived from Kasalath reduced 3.31% of the grain weight loss under heat stress. One located between R1613-C970 on chromosome 1 and the other between C1226-R1440 on chromosome 7, with additive effect 2.38 and 2.92%, respectively. The tolerance alleles of both these QTLs were derived from Nipponbare. Both of these QTLs had significant QTL×environment interactions, and the latter was involved in epistatic interaction also. Eight pairs of epistatic effect QTLs were detected, one pair each on chromosomes 1,2,3, 5, 7, 8, 10 and 12. The results could be useful for elucidating the genetic mechanism of heat-tolerance and the development of new rice varieties with heat tolerance during grain filling phase.     

Differential responses to high temperature during maturation in heat-stress-tolerant cultivars of Japonica rice

High-temperature stress during the grain-filling stage reduces grain quality of rice, and this is a serious problem in Japan, especially in the Kyushu region. To solve this problem, various heat-tolerant cultivars have been bred, such as ‘Nikomaru’, ‘Kumasannochikara’, ‘Genkitsukushi’, ‘Sagabiyori’, and ‘Otentosodachi’. When cultivated under high temperature after flowering, these heat-tolerant cultivars had lower percentages of chalky grains than in the heat-sensitive cultivar ‘Hinohikari’. All the heat-tolerant cultivars markedly decreased the nonstructural carbohydrate content in the stem under the high temperature compared to control condition during early grain-filling stage, which is considered to be a common trait of heat tolerance. Notably, ‘Sagabiyori’, ‘Genkitsukushi’, and ‘Nikomaru’ maintained a nucellar epidermis at 17 days after flowering (DAF) under high temperature, whereas the nucellar epidermis disappeared in ‘Hinohikari’. In addition, the expression of AGPS2b, thought to be a rate-limiting enzyme in starch synthesis, in ‘Kumasannochikara’, ‘Otentosodachi’, and ‘Nikomaru’ did not decrease under high temperature, whereas ‘Hinohikari’, ‘Sagabiyori’, and ‘Genkitsukushi’ could not maintain expression of the gene at 17 DAF. Moreover, the expression of Amy3E, a starch-degradation-related gene considered to induce grain chalkiness, in ‘Kumasannochikara’ at 17 DAF was not increased by high temperature. These results suggest that the heat-stress-tolerant cultivars have various mechanisms for dealing with high-temperature stress.     

水稻粒形遗传与长粒型优质粳稻育种进展

水稻粒形遗传机制和粒形育种一直是水稻品质和产量研究的重要领域。在粳稻品种选育中,对长粒形遗传种质的选择,已经成为近年来一种新的育种趋势。我们在总结粒长及其相关性状的遗传研究进展和已知粒长相关基因功能的基础上,系统地介绍了南方稻区长粒粳稻的育种历程及其选育策略。同时,对以品种嘉禾218为代表的南方长粒晚粳稻系列品种农艺性状和品质特性进行了比较,在阐明长粒粳稻优质、高产的品种特点的同时,提出长粒晚粳育种中选择粒形、兼顾株型、保产提质的选育思路。     

稻米垩白性状对高温耐性的QTL分析

【目的】本研究旨在筛选与稻米外观品质高温耐性连锁的分子标记,为稻米品质育种提供参考。【方法】以耐热水稻品系996和热敏感水稻品系4628为亲本构建的重组自交系为材料,采用垩白粒率耐热指数、垩白大小耐热指数和垩白度耐热指数为评价指标,对水稻垩白性状的高温耐性QTL进行检测。【结果】采用复合区间作图法两年共检测到垩白性状高温耐性QTL 24个,包括垩白粒率高温耐性QTL 8个,垩白大小高温耐性QTL 12个,垩白度高温耐性QTL 4个。其中,第6染色体上的2个垩白粒率高温耐性QTL和第7染色体上的2个垩白度高温耐性QTL在两年中重复检测到,且这2个稳定表达的垩白度位点与2015年检测到的第7染色体上的垩白粒率位点重合。另外,发现有4个QTL一因多效,同时影响垩白粒率、垩白大小及垩白度。【结论】控制垩白粒率耐热指数的q HTCGR6.1和控制垩白度耐热指数的q HTCD7.1是新的QTL。     

优质籼型不育系浙农8010A的选育与应用

不育系浙农 ８０１０ Ａ 育性败育彻底、开花习性好、异交率高、配合力强,尤其是其低直链淀粉含量对改善杂交籼稻的米质有广阔的应用前景。     

Responses of Yield Characteristics to High Temperature During Flowering Stage in Hybrid Rice Guodao 6

By sowing at different dates during 2005 and 2006 both in paddy fields and greenhouse, a super hybrid rice combination Guodao 6 and a conventional hybrid rice combination Xieyou 46 (as control) were used to analyze the differences in heat injury index, seed setting rate, grain yield and its components. Guodao 6 showed more stable yield and spikelet fertility, and lower heat injury index than Xieyou 46. Further studies indicated that the spikelet sterility is positively correlated with the average daily temperature and the maximum daily temperature, with the coefficients of 0.8604 and 0.9850 (P<0.05) respectively in Guodao 6. The effect of high temperature injury on seed setting caused by maximum daily temperature was lower than that by average daily temperature during the grain filling stage.     

水稻着粒密度的遗传分析

 水稻小穗着粒密度是一个重要的农艺性状,其与单株穗数、小穗数和粒重等有密切的联系。该性状在密穗品种与疏穗品种杂交的 F_1值高于双亲平均值,F_2群体分布呈近正态分布的曲线。因此该性状属多基因制约的数量性状,而且和单株生产力有显著相关,故必将受到水稻育种家的重视。