Regulation of Spikelet Development in Rice by Hormones

Four indica rice varieties belonging to two maturity groupswere cultivated in medium irrigated land during the wet seasonof 1989. In all varieties, the primary branches of the panicledeveloped basipetally. Poor development of spikelets on thebasal branches resulted in a high percentage (50–60%)of degeneration and sterility and, consequently, grain yieldwas poor. Application of gibberellic acid (GA3) and kinetin(6-furfuryl amino purine) improved development and grain yieldon all branches. The treatments were more effective on the proximalbranches and reduced the existing gradient in growth, developmentand yield between the distal and proximal branches. In contrastto this, 3-indole acetic acid (IAA) treatment increased spikeletgrowth and development in the distal branches, but suppressedthem in the proximal branches thus increasing the heterogeneityin primary branch growth and development. The soluble carbohydrateand amino acid content per unit dry weight of the primary branchesincreased at the time of anthesis following the applicationof growth regulators. The assimilate level declined rapidlyat the post-anthesis stage. Associated with the depressed growthand development in the lower branches of the panicle by IAAtreatment, was a significantly higher level of assimilates inthese branches in the post-anthesis period. The physiologicalmechanism of spikelet growth regulation is discussed. Key words: Rice, assimilates, development, spikelets, fertility, hormones     

Apoplasmic assimilates and grain growth of contrasting rice cultivars differing in grain dry mass and size

Apical dominance in assimilate filling impacts grain growth in basal spikelets of rice panicle. In this study, organic materials of the pericarp, apoplasmic space and endosperm of the apical and basal caryopses, and photosynthesis of the flag leaf were measured during early part of grain development in three types of rice cultivars with similar phenology, but difference in grain weight and size in the dry and wet seasons of 2006 and 2007, respectively. Photosynthetic activity of the flag leaf was consistently low in small-seeded cultivars. Rates of grain filling and cell division of endosperm and concentration of assimilates, starch, proteins and chlorophylls of the caryopsis were lower, but spikelet ethylene production and peroxidase activity were higher in a small-seeded cultivar compared to a big-seeded cultivar. Similar disparities in grain filling and other attributes were noticed for the inferior basal spikelets of the panicle compared to the superior apical spikelets, except the assimilate concentration of the pericarp and endosperm. Temporal fluctuation in assimilate concentration of the organs were similar between the cultivars. Concentration of apoplasmic assimilates mostly exhibited negative correlation with that of pericarp and endosperm. Compared to the apical spikelets, correlation was more negative for the basal spikelets. Conversely, correlation was positive between the concentration of apoplasmic assimilates and endosperm cell number and grain weight of the cultivars. Ethylene released from the spikelets at anthesis affected growth and cell division rates of endosperm and enhanced protein and chlorophyll degradation and peroxidase activity of the caryopsis. It was concluded that variation in spikelet ethylene production may be responsible for differences in size or weight of grains among rice cultivars and spikelets at different locations of the panicle. The concentration of apoplasmic assimilates could be an indicator for grain filling capacity, and ethylene regulated the concentration by affecting pericarp activity for assimilate unloading.     

Apical control of branch growth and angle in woody plants

Apical control is the inhibition of a lateral branch growth by shoots above it (distal shoots). If the distal shoots are cut off to remove apical control, the lateral branch can grow larger and may bend upwards. Apical control starts when new lateral buds grow after passing through a period of dormancy. Buds initially break and produce leaves, then apical control is exerted and the lower (proximal) laterals stop growing. Apical control also inhibits growth of large, old branches. Gravimorphism and restricted water and nutrient transport can inhibit branch growth, but they are not primary mechanisms of apical control. Apical control may reduce branch photosynthesis. Under apical control allocation of branch-produced assimilate to the stem is relatively high, so low assimilates in the branch may limit branch growth even though hormone levels are adequate for growth. Hormones appear to be involved in apical control, but it is not known how. One role of hormones may be to maintain the strength of the stem sink for branch-produced assimilate. Upward bending of a woody branch after release from apical control requires both new wood production and production of wood cells that can generate an upward bending moment. Apical control inhibits radial growth of branches and, in some species, may regulate the production of wood with an upward bending moment.     

近地层臭氧浓度升高对杂交稻颖花形成的影响

依托全球唯一的稻田开放式空气中臭氧浓度增高系统平台,以汕优63和两优培九为供试材料,设置大气背景臭氧浓度和高臭氧浓度（比大气背景臭氧浓度高50%）两个浓度水平,研究FACE条件下高O₃浓度对杂交稻颖花形成的影响.结果表明：高O₃浓度使汕优63和两优培九每穗颖花数分别减少28朵和34朵,下降幅度分别为15%和13%.从稻穗构成看,高O₃浓度胁迫下杂交稻每穗颖花数减少主要与每穗2次枝梗颖花数明显减少有关,对每穗1次枝梗颖花数的影响较小,因此高O₃浓度胁迫下水稻每穗1次枝梗颖花数占全穗的比率增加,每穗2次枝梗颖花数占全穗的比率降低.从颖花形成看,高O₃浓度胁迫下杂交稻每穗颖花数下降主要是颖花（特别是2次颖花）的分化受到抑制所致,而颖花的退化数不增反降.上述结果表明,采取相应措施削弱高O₃浓度胁迫对颖花分化的抑制作用可能是近地层高O₃浓度条件下减少杂交稻产量损失的关键.     

异季条件下稻穗枝梗和颖花分化、退化及其形成的差异性研究

以19个水稻杂交组合为试验材料,通过大田条件下的分期播种,于抽穗期调查了稻穗枝梗、颖花分化、退化及形成有关性状,分析了各性状不同季节的变异及其相互关系。结果表明,单穗一次枝梗分化数、二次枝梗分化数、颖花分化数、一次枝梗退化数、二次枝梗退化数、颖花退化数、一次枝梗退化率、二次枝梗退化率、颖花退化率、一次枝梗形成数、二次枝梗形成数、一次枝梗颖花形成数、二次枝梗颖花形成数和单穗颖花总形成数等性状均存在较大的组合间差异和季节间变异,其中19个组合单穗颖花总形成数异季差异率变幅为5.12%~93.59%,平均变异率为34.19%;总体上,不同季节下二次枝梗各性状的变异性强,颖花性状其次,一次枝梗各性状变异性最小;根据异季条件下稻穗枝梗和颖花分化、退化及其形成性状的变异性,从大穗角度讨论了杂交稻新组合育种资源选择与利用问题。     

水稻穗型突变体cl的鉴定、保护与遗传分析

通过神州八号飞船搭载广东省常规优质稻主导品种粤农丝苗获得一个稳定遗传的穗型突变体cl,中文名为粤花占1号,2015年完成农业部植物新品种权的申请工作(申请号:20151728.8,公告号:CNA014491E)。该突变体农艺性状与野生型极为相似,熟色好,产量高,但表现出异常的花序结构,一次枝梗数增加,二次枝梗、小穗梗长度严重缩短,枝梗顶端小穗3粒簇生在一起。进一步的细胞学观测发现,差异主要源自幼穗分化的第3期到第4期发育异常所致。遗传分析表明该性状由非完全显性单基因控制,BSA性状定位法初步将该基因定位于染色体6,命名为CL-6。本研究的结果为进一步精细定位和克隆CL-6基因、找到控制水稻幼穗枝梗伸长缩短的功能基因或调控因子奠定基础,同时为培育理想穗型水稻和观赏农业提供优良种质。     

地方稻资源D43的开花期耐热特性研究

水稻在抽穗开花期对高温胁迫非常敏感,通过挖掘耐热资源,培育耐热水稻品种是应对高温热害最有效的方式。前期研究发现,地方稻资源D43在花期连续高温条件下能保持较高的结实率。本研究在大田和人工气候室不同高温处理下,分析了D43的开花时间与耐热性之间的相互关系。结果表明,高温能够使水稻的开花时间提前,D43表现出稳定的早花时特性,高温胁迫下的开花时间集中在8∶30~10∶00;在开花时间段恒定高温胁迫下,D43的结实率较低;但在大田高温和人工气候室模拟高温胁迫下,D43的开花时间避开了日高温段,从而表现出较高的结实率;花器官形态性状包括花药开裂率、柱头上的花粉附着数、花粉萌发数与结实率之间呈显著正相关,因此可用于评价水稻的花期耐热性。     

Inflorescence, spikelet, and floral development in Panicum maximum and Urochloa plantaginea (Poaceae)

Inflorescence development in Panicum maximum and Urochloa plantaginea was comparatively studied with scanning electron and light microscopy to test the transfer of P. maximum to Urochloa and to look for developmental features applicable to future cladistic studies of the phosphoenol pyruvate carboxykinase (PCK) subtype of C(4) photosynthesis clade (P. maximum and some species of Brachiaria, Chaetium, Eriochloa, Melinis, and Urochloa). Eleven developmental features not discernable in the mature inflorescence were found: direction of branch differentiation; origins of primary branches; apical vs. intercalary development of the main axis; direction of spikelet differentiation; direction of glume, lemma and palea differentiation; position of the lower glume (in some cases); size of the floret meristem; pattern of distal floret development; pattern of gynoecium abortion; differential pollen development between proximal and distal floret; and glume elongation. Inflorescence homologies between P. maximum and U. plantaginea are also clarified. Panicum maximum and U. plantaginea differ not only in their mature inflorescence structure but also in eight fundamental developmental features that exclude P. maximum from Urochloa. The following developmental events are related to sex expression: size of floret meristem, gynoecium abortion, pollen development delay in the proximal floret, glume elongation and basipetal floret maturation at anthesis.     

<Emphasis Type="Italic">OsREL2</Emphasis>, a rice TOPLESS homolog functions in axillary meristem development in rice inflorescence

The morphology of the rice inflorescence, called the panicle, is determined mainly by the activities of axillary meristems including primary, secondary, and spikelet meristems. Recently, in maize, the RAMOSA1 ENHANCER LOCUS2 (REL2) gene, orthologous to the Arabidopsis shoot apical meristem fate-determining TOPLESS, was shown to be involved in the regulation of axillary meristem determinacy. In order to investigate the function of the rice REL2 homolog, we identified and characterized the rice REL2 gene (OsREL2). Compared to other rice TPL homologs, OsREL2 gene expression stayed relatively low throughout panicle development. We characterized a T-DNA insertion osrel2 mutant that showed pleiotropic phenotypic defects, such as defects in panicle heading, sterile lemma elongation, and panicle development, suggesting the OsREL2 functions in multiple developmental processes. In particular, osrel2 developed shorter axillary branches and reduced numbers of lateral organs on axillary branches in comparison to the wild-type, indicating that OsREL2 is important in axillary meristem maintenance. Interestingly, osrel2 produced more primary branches and fewer secondary branches than the wild-type. These results suggest that OsREL2 is involved in branch formation regulation, presumably by suppressing primary branch formation and promoting secondary branch formation.     

水稻穗型的遗传调控研究进展

穗型作为水稻(Oryza sativa)重要的农艺性状,近年来一直是研究热点。该文简要介绍了水稻穗部发育的一般过程,总结了近年来发现的调控水稻穗型相关基因,并根据水稻幼穗发育过程将其分为4类:分别调控枝梗分生组织的形成、枝梗分生组织的大小、小穗分生组织的转变时间以及枝梗的伸长;并概括分析了上述基因在调控水稻幼穗发育过程中所呈现出的路径关系。最后对水稻穗型遗传调控研究的未来发展方向进行了展望。     

基因型×环境互作效应对水稻穗部性状杂种优势的影响

采用包括基因型×环境互作效应的加性显性加性×加性上位性遗传模型,分析了籼粳杂交穗部性状杂种优势的两年资料。结果表明,7个穗部性状的杂种优势既由基因型控制也受环境(年份)互作效应影响。主穗粒数、主穗长、一次和二次枝梗总长和着粒密度表现较强的正向群体平均优势,而一次和二次枝梗数则表现显着的负向优势。基因型×环境互作预测结果表明,主穗粒数、主穗长、一次枝梗总长在两年中的环境互作表现正效应,一次和二次枝梗数则为负效应,着粒密度和二次枝梗总长表现不同方向的效应。利用IR66158-37(P₁)、IR65600-85(P₂)、明恢63(P₄)和R669(P₆)进行杂交配组,其杂种后代易获得重穗型的穗部结构,可在超高产育种中加以利用。     

Effect of awns and drought on the supply of photosynthate and its distribution within wheat ears

The presence of awns doubled the net photosynthetic rate of wheat ears and also increased the proportion of ¹⁴CO₂ assimilated by the ear that moved to the grain. The effect of water supply on photosynthesis and movement of assimilates was greater for leaves than ears, so that drought increased the proportion of assimilate contributed by ear photosynthesis to grain filling from 13% to 24% in the awnless ears, and from 34% to 43% in the awned ears. ¹⁴C assimilated by the ears was most important to the economy of the upper spikelets and to the distal florets in each spikelet, whereas flag leaf assimilate went mainly to the spikelets in the lower half of the ear, and to the proximal florets. Awns increased grain yield in the dry but not in the irrigated treatment, despite the large contribution of awned ears to grain filling. Either the supply of assimilate did not limit grain yield when water supply was not limiting, or there were compensating disadvantages to awns. However, they did not seem to have any adverse effect on the development of the upper florets, nor did they reduce grain number per ear.     

Glutelin accumulation and changes in the levels of its mRNA in the superior and inferior spikelets of rice ear during ripening

Glutelin accumulation in the apical spikelet of the top primary branch (superior spikelet) and the second spikelet of the lowest secondary branch (inferior spikelet) of the ear of the rice plant (Oryza sativa L.) was characterized during grain filling.In the superior spikelet, the accumulation of dry matter and nitrogen started immediately after flowering and rapidly reached the maturation level by 20 days after heading (DAH). At 7 DAH, total RNA content had already reached its maximum level and glutelin mRNA content 70% of its maximum. The increase in glutelin mRNA was followed by a rapid increase in glutelin between 7 and 16 DAH.In the inferior spikelet dry matter, nitrogen and glutelin accumulation were low immediately after flowering and increased only after grain filling of the superior spikelet was almost complete. Total RNA and glutelin mRNA increased much later at slower rates than in the superior spikelet.It is very likely that the retardation of dry matter, total nitrogen and glutelin accumulation in the inferior spikelet is due to retardation of differentiation and development of endosperm tissue, and to glutelin gene expression in endosperm cells. It is suggested that the delayed development resulted from limited partitioning of nutrients to the inferior spikelet at the early stage of ripening.     

不同穗型小麦光能利用和旗叶13C同化物分配特性及对补灌水平的响应

为明确不同穗型小麦冠层光能利用和¹³C同化物分配特性的差异及对补灌水平的响应,以中穗型品种‘青农2号’和‘济麦22’、大穗型品种‘山农23’和‘山农30’为材料,设置3个水分处理:小麦全生育期不灌水(W₀)、节水灌溉(W₁,拔节期和开花期0～40 cm土层土壤相对含水量分别补灌至65%和70%)和充分灌溉(W₂,拔节期和开花期0～40 cm土层土壤相对含水量分别补灌至85%和90%),研究了不同处理对两种穗型小麦冠层光能利用和旗叶¹³C同化物分配特性的影响.结果表明:W₁处理两种穗型小麦品种开花后2、11、20和31 d的叶面积指数、冠层PAR截获率和利用率均显著高于W₀处理,再增加灌水至W₂处理,上述各指标无显著变化.¹³C示踪表明,济麦22和山农23的W₁旗叶¹³C同化物在籽粒中的分配量比W₀分别高159.34和171.1 g·hm^-2,分配比例分别高6.5%和6.5%,与W₂无显著差异;两种穗型品种W₁的籽粒产量亦显著高于W₀,与W₂无显著差异.不同穗型品种比较,节水灌溉条件下中穗型品种在开花后2和11 d、大穗型品种在开花后20和31 d具有较高的冠层光合有效辐射(PAR)截获和利用能力;中穗型品种济麦22旗叶¹³C同化物在籽粒的分配量和分配比例比大穗型品种山农23分别低6.8%和2.7%.     

不同水稻品种对三化螟抗性差异的机理

研究了不同水稻品种对三化螟Scirpophaga incertulas抗性差异的机理。结果表明, 在汕优63、镇稻2号和92-109稻株中, 虫体的游离必需氨基酸含量极低（0.422～0.875 nmol/mg）, 幼虫生长发育缓慢;而在9-92、武育粳3号等稻株中, 虫体的游离必需氨基酸含量高（2.788～5.421 nmol/mg）, 幼虫生长快。虫体的游离必需氨基酸含量和生长发育的差异, 与不同水稻品种的游离必需氨基酸含量、总必需氨基酸含量、还原糖含量和叶鞘硅化细胞密度均无明显相关。叶鞘维管束间距、鞘脊宽度及小于蚁螟头宽的叶鞘维管束间距百分率的差异, 是不同水稻品种抗螟性差异的主要作用因子。     

Genetic control of branching patterns in grass inflorescences

Inflorescence branching in the grasses controls the number of florets and hence the number of seeds. Recent data on the underlying genetics come primarily from rice and maize, although new data are accumulating in other systems as well. This review focuses on a window in developmental time from the production of primary branches by the inflorescence meristem through to the production of glumes, which indicate the transition to producing a spikelet. Several major developmental regulatory modules appear to be conserved among most or all grasses. Placement and development of primary branches are controlled by conserved auxin regulatory genes. Subtending bracts are repressed by a network including TASSELSHEATH4, and axillary branch meristems are regulated largely by signaling centers that are adjacent to but not within the meristems themselves. Gradients of SQUAMOSA-PROMOTER BINDING-like and APETALA2-like proteins and their microRNA regulators extend along the inflorescence axis and the branches, governing the transition from production of branches to production of spikelets. The relative speed of this transition determines the extent of secondary and higher order branching. This inflorescence regulatory network is modified within individual species, particularly as regards formation of secondary branches. Differences between species are caused both by modifications of gene expression and regulators and by presence or absence of critical genes. The unified networks described here may provide tools for investigating orphan crops and grasses other than the well-studied maize and rice.

Recent work on grass inflorescence branching identifies extensive conserved regulation, but also divergence particularly in formation of secondary branches and spikelets.     

Quantitative trait locus analyses of ozone-induced grain yield reduction in rice

Reduction of grain yield (total seed weight) by ozone in rice (Oryza sativa L.) is believed to be caused by ozone-induced reduction of photosynthetic activity followed by growth inhibition. Here, japonica rice cultivar Sasanishiki and indica rice cultivar Habataki showed different responses to ozone. When exposed to ozone, the leaves of Habataki exhibited no critical damage, whereas those of Sasanishiki developed lesions. Conversely, ozone exposure reduced total seed weight by 19% in Habataki, but not significantly in Sasanishiki. Chronic ozone exposure also significantly decreased culm length, number of primary rachis branch, and number of spikelets per panicle in Habataki. QTL analysis in Sasanishiki/Habataki chromosome segment substitution lines identified a single locus associated with the yield loss caused by ozone on chromosome 6 of Habataki close to marker RM3430 (107.6 cM). A QTL for reduction of primary rachis branch number and total spikelet number was found in the same position. These results indicate that a QTL on chromosome 6 has an important role in ozone-induced yield loss, and is also involved in primary rachis branch formation and total spikelet number in ozone-exposed rice.     

高温下干旱和渍水对冬小麦花后旗叶光合特性和物质转运的影响

以蛋白质含量不同的两个冬小麦品种扬麦9号和豫麦34为材料,研究了不同温度和水分条件下小麦花后旗叶光合特性的变化、营养器官花前贮藏干物质和氮素转运特征及其与籽粒产量和品质形成的关系.结果表明,高温及干旱和渍水均明显降低了旗叶光合速率和叶绿素含量（SPAD值）,但高温下干旱和渍水对光合作用的影响加重.小麦营养器官花前贮藏干物质、氮素转运量和转运率在适温下表现为干旱>对照>渍水,高温下则表现为对照>干旱>渍水.适温下花后同化物积累量表现为对照>渍水>干旱,高温下则表现为对照>干旱>渍水.花后氮素积累量在适温和高温下均表现为对照>渍水>干旱.籽粒淀粉含量以适温适宜水分处理最高,高温渍水下最低;蛋白质含量以高温干旱下最高,适温渍水下最低.温度和水分逆境下小麦粒质量和淀粉含量的降低与花后较低的光合能力及干物质积累有关,而蛋白质含量则与花前贮藏氮素的转运量和转运率有关.     

Assimilatverteilung in Sommergerstensorten mit unterschiedlicher Resistenz gegenüber dem Echten Mehltau (Erysiphe graminis f. sp. hordei)

Distribution of assimilates in cultivars of spring barley with different resistance against powdery mildew (Erysiphe graminis f. sp. hordei) Transport and distribution of radioactive labelled assimilates in spring barley cultivars with different degrees of resistance to powdery mildew were studied after ¹⁴CO₂-treatment of single leaves. Plants of the cultivars ‘Amsel’ (susceptible), ‘Asse’ (adult plant resistant), and ‘Rupee’ (resistant) were analyzed at the vegetative growth stage (5. leaf unfolded) and the generative growth stage (anthesis). At the vegetative growth stage the assimilate export from the mildew inoculated 5. leaf of ‘Amsel’ and ‘Rupee’ is decreased; in ‘Asse’, there is no considerable change of assimilate distribution due to infection. At the generative growth stage the assimilate export from the infected flag leaf of ‘Amsel’ is reduced when the fungus, is sporulating. In the cultivar ‘Asse’ the assimilates are bound at the infection site until the seventh day after inoculation, then the transport of assimilates to the ear is increased. In ‘Rupee’ mildew inoculation causes an enhanced assimilate transport to the ear. The changes in assimilate distribution due to mildew inoculation are discussed with respect to the different types of host-parasite-interactions and the source-sink-activities in the different cultivars.