Spikelet Sterility and Flowering Response of Rice to Water Stress at Anthesis

Water deficits at the anthesis stage of rice (Oryza sativa L.)induce a high percentage of spikelet sterility and reduce grainyield. This study attempted to elucidate the direct effectsof water stress on panicle exsertion, spikelet opening, andspikelet desiccation leading to spikelet sterility. A well-wateredtreatment and two water stress levels were imposed in pot-grownplants of IRAT 13 (upland cultivar) and IR20 (lowland cultivar)at the time of flowering under greenhouse conditions A cultivar difference was observed in the flowering responseto water stress with a high sensitivity in IR20. The time courseof panicle exsertion showed an inhibitory effect due to thelow panicle water status. Low panicle water potentials significantlyreduced the number of opened spikelets. Spikelet opening wascompletely inhibited at panicle water potentials below –1·8MPa and –2·3 MPa in IR20 and IRAT 13, respectively.However, the peak spikelet opening time in a day was not influencedby the stress treatment. Spikelets in stressed panicles wereobserved to remain open for a longer period than in the well-wateredpanicles. The role of turgor in spikelet opening is also discussedin the study. At low panicle water potentials, severe desiccationof spikelets and anthers was noted. The deleterious effectsof water deficits on spikelet opening and spikelet water losscontributed to reduced spikelet fertility Oryza sativa L., rice, spikelet sterility, flowering, water stress, panicle water potentials, turgor potentials, desiccation     

Abscisic Acid and Ethylene Interact in Rice Spikelets in Response to Water Stress During Meiosis

This study investigated whether and how the interaction between abscisic acid (ABA) and ethylene is involved in the regulation of rice (Oryza sativa L.) spikelet sterility when subjected to water stress during meiosis. Two rice cultivars, HA-3 (drought-resistant) and WY-7 (drought-susceptible), were used and subjected to well-watered and water-stressed (WS) treatments during meiosis (15–2 days before heading). Leaf water potentials of both cultivars markedly decreased during the day as a result of the WS treatment, but panicle water potentials remained constant. The percentage of sterile spikelets in WS plants was increased by 49.7% for WJ-7 but only 12.7% for HA-3. ABA, ethylene, and 1-aminocyclopropane-1-carboxylic acid were all enhanced in spikelets by the water stress, but ethylene was enhanced more than ABA in WY-7 when compared with that in HA-3. Spikelet sterility was significantly reduced when ABA or amino-ethoxyvinylglycine, an inhibitor of ethylene synthesis, was applied to the panicles of WS plants at the early meiosis stage. Application of ethephon, an ethylene-releasing agent, or fluridone, an inhibitor of ABA synthesis, had the opposite effect, and sterility was increased. The results suggest that antagonistic interactions between ABA and ethylene may be involved in mediating the effect of water stress on spikelet fertility. A higher ratio of ABA to ethylene would be a physiologic trait of rice adaptation to water stress.     

Comparative Proteomic Analysis Reveals Nitrogen Fertilizer Increases Spikelet Number per Panicle in Rice by Repressing Protein Degradation and 14-3-3 Proteins

The spikelet number per panicle is established in the early stages of panicle development. Nitrogen fertilizer application before panicle initiation is known to increase spikelet number, which is one of the most important traits in rice productivity determination. However, the basic proteomic mechanism remains poorly understood. The present study shows that nitrogen fertilizer significantly increased spikelet number and grain yield in rice. Proteomic variations were further analyzed in young panicles at the secondary panicle branch initiation and spikelet meristem initiation under nitrogen fertilizer treatment. Proteomic analysis identified 63 proteins with significant differential accumulation in young panicles under nitrogen fertilizer treatment. Proteolysis represents the largest functional category, which suggests that protein degradation is an important pathway in the response to nitrogen fertilizer. Importantly, nitrogen fertilizer significantly reduced 14-3-3 proteins, which interact with key enzymes associated with carbon and nitrogen metabolism, and the rice FT homologue Hd3a. Real-time PCR revealed that Hd3a signaling is also repressed by nitrogen fertilizer in leaves. This study contributes to a better understanding of the regulation of nitrogen fertilizers in the flowering pathway leading to panicle development. The identification of novel genes provides new insight into the profound impacts of nitrogen fertilizer on panicle development in rice.     

Relationship Between Male Fertility and Endogenous Phytohormones in Photoperiod-Sensitive Genic Male-Sterile Rice

A spontaneous male sterile rice plant (Oryza sativa L. cv. Nongken 58S) "Photoperiod-sensitive Genic Male-sterile Rice has been found to be male sterile under long day cycles (LD) and fertile in short day cycles (SD). The period from secondary rachis-branch and spikelet primodia to pollen mother cell formation in the process of panical development was the photoperiod-sensitive stage for fertility alternation. The phenotype of this mutant was reported to be controlled by two pairs of recessive alleles. The research on relationship between the fertility alternation and phytohormone action in this mutant have been performed by Chinese scientists since 1985. In order to study the mechanism of fertility alternation in Nongken 58S, endogenous IAA, ABA, GA1 and GA4 in apical leaves and reproductive organs in different development stages under LD and SD conditions have been quantiatvely and qualitatively identified by GC-MS-SIM method. It was found that endogenous IAA in apical leaves at the stage of pistil and stamen primodia formation and in panicles at pollen mother cell stage of Nongken 58S with LD condition was deficient comparing with those in SD. Endogenous ABA level in panicles at pollen mother cell stage, in spikelets at uninucleate stage and in anthers at anthesis stage of Nongken 58S-LD were lower than those in SD. ABA levels in corresponding organs and developmental stages of wild type of rice, "Nongken 58" were always higher in LD treatment than those in SD. Endogenous IAA, GA1 and GA4 levels in anthers at anthesis stasge of "Nongken 58"-LD were increased obviously. Thus it appeared that "Nongken 58" possess stronger resistance to LD stress than Nongken 58S. It is concluded that IAA deficiency of reproductive organs at early developmental stage, ABA decrease implying poor resistance to LD stress and reduction of GAs in late developmental stages were the factors causing the anther sterility in Nongken 58S-LD.     

Drought-induced spikelet sterility is associated with an inefficient antioxidant defence in rice panicles

Water stress-induced spikelet sterility limits rice production under upland conditions. The causes of spikelet sterility under drought stress are poorly understood. In this study the role of antioxidant defence management in drought-induced spikelet sterility was investigated in two rice ( Oryza sativa ) genotypes differing in drought resistance. Drought-resistant N22 genotype showed less water stress-induced spikelet sterility when compared to the susceptible N118 genotype under upland conditions. The N22 panicles maintained higher RWC and turgor potential and lower H₂O₂ levels across the developmental stages under water stress than that of N118 panicles. Drought-induced enhancement in superoxide dismutase (SOD, EC 1.15.1.1) activity coupled with higher ascorbate (AsA), glutathione (GSH) content and enhanced ascorbate peroxidase (APX, EC 1.11.1.11) and glutathione reductase (GR, EC 1.6.4.2) activities resulted in lower H₂O₂ levels in N22 panicles. In contrast, insufficient enhancement in SOD, APX and GR activities resulted in relatively higher H₂O₂ levels under water stress in N118 panicles. The N22 panicles exhibited a higher number of SOD and APX isozymes in comparison with N118 panicles that might provide better reactive oxygen species scavenging. Hence it is concluded that well-equipped antioxidant defence plays an important role in minimizing water stress-induced spikelet sterility in upland rice.     

不同生育阶段夜温升高对双季水稻产量的影响

利用两间玻璃温室内夜间不同的温度条件,研究了水稻不同生育阶段的夜温升高对双季早、晚稻产量的影响.结果表明：播种-幼穗分化（一次枝梗分化期）期间夜温升高,有利于双季水稻的分蘖,植株有效穗数显著增加,平均夜间最低温度每升高1 ℃,双季早、晚稻的产量分别提高10.02%～13.18%和6.52%～7.78%（P＜0.01）;幼穗分化-抽穗（10%稻穗抽出剑叶鞘）期间夜温升高,导致颖花退化,使每穗发育颖花数减少,平均夜间最低温度每升高1 ℃,双季早、晚稻的产量分别下降3.76%～6.67%和3.66%～6.94%（P＜0.01）;灌浆结实期（抽穗-成熟）夜温升高,双季早稻的结实率和产量显著下降,而双季晚稻的结实率和产量显著提高,平均夜间最低温度每升高1 ℃,双季早稻的产量下降2.07%～5.61%（P＜0.05）,双季晚稻的产量提高1.63%～2.28%（P＜0.05）.表明不同生育阶段的夜温升高对双季水稻产量的影响存在明显差异.     

云南不同类型一季稻产量形成及其与气象因子的关系

利用云南省14个不同海拔农业气象观测站1994-2010年水稻大田发育期和产量观测资料,研究水稻产量形成及气象因子对低产水稻产量构成因素的影响;根据14个站点水稻产量构成因素的系统聚类分析结果和水稻类型,将水稻分为低产粳型、低产籼型、高产粳型和高产籼型4个类型.对这4类水稻产量构成因素的分析结果表明,单位面积颖花量与产量呈极显著正相关,低产粳型水稻产量主要受结实率和颖花量影响,其他3类颖花量对产量影响较大.低产粳型水稻主要受低温的影响,孕穗期低温降低颖花量和有效穗数,孕穗期和抽穗开花期低温增加空壳率,以平均气温、平均最高气温和冷积温的影响较大;乳熟前期较低的平均气温增加秕谷率,同时降低千粒重.低产籼型水稻产量构成因素受多种气象因子的综合影响明显;分蘖期和拔节期一定程度的增温不利于增加低产籼型水稻有效穗数,而分蘖期较多的日照时数和较大的平均气温日较差有利于有效穗数的增加,分蘖期和拔节期气温和日照时数与每穗粒数的关系呈“抛物线型”;低产籼型水稻空壳率一定程度上受抽穗开花期低温的影响,而乳熟前期高温少雨不仅增加秕谷率,还导致千粒重明显下降.     

光温条件对短光低温不育水稻育性转换的影响

短光低温不育水稻是一类与长光高温不育水稻育性转换特性相反的新种质,通过对分期播种材料自然条件下的育性及人工短光处理后的育性比较分析表明,宜DIS的光敏期为第一苞分化期至第二次枝梗原基和小穗原基分化期（幼穗分化I－III期）,在常温下光敏期的短日处理可使育性趋于不育,但在高温下光敏期短日不育效应被削弱,宜DIS的花粉育性在高,低温作用下育性的均出现下降,根据高,低温影响育性的资料分析得到的敏感期基本一致,即花粉母细胞形成期至花粉内容物充实期（幼穗分化V－Ⅶ期）。     

Influence of time and level of nitrogen application of growth and yield of rice

Summary Results of a greenhouse sand-culture experiment conducted to study the effect of increasing or decreasing nitrogen supply at various stages of growth on the growth, yield, and yield components of rice are reported. Adequate nitrogen during the vegetative phase of growth encouraged tillering which had a bearing on the number of panicles per plant. Excess nitrogen during the reproductive phase of growth produced panicles with unfilled or partly filled grains. Similar effects were observed when the nitrogen status of the growth medium was changed from a low to a high level at the stages of reduction division and anthesis. Decrease in yield was observed when plants growing at a low level of nitrogen were transferred to a high level at the end of the vegetative phase. High levels of nitrogen in the growth medium at time of heading and after increased the concentration of nitrogen in the grain. There was an inverse relationship between yield of filled grain and nitrogen in the grain.     

Pollen germination and in vivo fertilization in response to high‐temperature during flowering in hybrid and inbred rice

High‐temperature during flowering in rice causes spikelet sterility and is a major threat to rice productivity in tropical and subtropical regions, where hybrid rice development is increasingly contributing to sustain food security. However, the sensitivity of hybrids to increasing temperature and physiological responses in terms of dynamic fertilization processes is unknown. To address these questions, several promising hybrids and inbreds were exposed to control temperature and high day‐time temperature (HDT) in Experiment 1, and hybrids having contrasting heat tolerance were selected for Experiment 2 for further physiological investigation under HDT and high‐night‐time‐temperature treatments. The day‐time temperature played a dominant role in determining spikelet fertility compared with the night‐time temperature. HDT significantly induced spikelet sterility in tested hybrids, and hybrids had higher heat susceptibility than the high‐yielding inbred varieties. Poor pollen germination was strongly associated with sterility under high‐temperature. Our novel observations capturing the series of dynamic fertilization processes demonstrated that pollen tubes not reaching the viable embryo sac was the major cause for spikelet sterility under heat exposure. Our findings highlight the urgent need to improve heat tolerance in hybrids and incorporating early‐morning flowering as a promising trait for mitigating HDT stress impact at flowering.     

Effect of sucrose level on rice grain deposition in detached panicle system

In detached IR36 rice panicles incubated in liquid medium for 10 days in 1.0–2.0% sucrose and 0.6% glutamine, final panicle weight increased but percentage and weight of grain protein decreased with 1.0–1.75% sucrose. Soluble sugars increased in stem and hull but not in developing grain. With five other rice panicles and IR36 panicles differing in grain size incubated in liquid culture containing 0.075% glutamine for 7 days, percentage grain protein was again lower in 1.5% sucrose than in 1.0% sucrose, with correspondingly heavier grain weight in four cases. Free-sugar levels of developing grains were lower in detached panicles than in the field grain samples in both experiments. Thus, sucrose level has a depressing effect on protein accumulation in the developing rice grain. Lysine content of grain protein decreased with increase in protein content.     

Effect of Growth Regulators and Chemicals on Pollen Sterility in TGMS Lines of Rice

Thermosensitive genic male sterility (TGMS) in rice is a widely adopted technique for successful hybrid rice production in Asia. TGMS lines remain male sterile when daily mean temperature is above the critical sterility temperature and are therefore used as female parents. The same line will remain fertile when mean temperature is below the critical sterility temperature. Achievement of 100% male sterility in TGMS lines is important for the successful utilization of TGMS lines as female parents in hybrid rice production. This study examined the external application of some growth regulators and chemicals and their effect on pollen sterility. Among the various treatments, ethrel (800 ppm), salicylic acid (600 ppm) and maleic hydrazide (0.2%) induced a significantly higher percentage of male sterility in the TGMS lines. The sprayed plants also showed higher total phenol accumulation in their flag leaves. The results suggest that it is possible to achieve 100% male sterility in TGMS lines with the external application of growth regulators and chemicals.     

Studies on the Physiology of Flowering of Timothy (Phleutm pratense L.): I. Influence of Daylength and Temperature on Initiation and Differentiation of the Inflorescence

Spikelet initiation is advanced and the proportion of plantswhich attain the reproductive condition is increased in S. 48timothy by lengthening the photo-period from 14 to 24 hours.In shorter periods of light, reproduction is almost completelyinhibited, and in 8-hour short days plants remain vegetativeeven after 35 weeks. Spikelet initiation at the shoot apex occursafter exposure to 3–5 long days followed by short days.Initiation also occurs when extended daylength is replaced by‘light-breaks’ during long nights, or when a singleleaf is photo-induced while the remainder of the plant receivesshort days. High temperatures promote spikelet initiation incontinuous light; in photoperiods nearer the threshold for floweringthis response is reversed and a rise in temperature from 55°to 75° F. increasingly inhibits reproduction. Once initiationhas occurred, spike differentiation is hastened by increasesin temperature or photoperiod. Internode elongation begins atthe time of spikelet initiation, and is promoted by temperatureand photoperiod. Elongated vegetative shoots may be producedwhen spikelet initiation fails in threshold photoperiods orhigh temperatures.     

Effect of Water Deficit Stress on Diffusive Resistance, Transpiration, and Spikelet Desiccation of Rice (Oryza sativa L.)

To understand the nature of panicle resistance to water loss,diffusive resistance, transpiration rate, and degree of tissuedesiccation were studied as affected by low tissue water potentialsand relative water contents (RWC) induced by water deficit stresses.A drought susceptible cultivar (IR20) and a moderately resistantcultivar (IRAT 13) were compared under moderate and severe waterdeficit stresses under greenhouse conditions. Irrigation waswithdrawn at flowering stage. During the 10-d stress period,changes in panicle and flag leaf transpiration and white headformation (desiccated) were monitored. Using cut panicles transpirationand white head formation at different RWC levels in 11 differentcultivars were also measured. In flag leaves, increased diffusiveresistance and leaf rolling reduced transpirational losses fromthe adaxial surfaces. Leaf resistance and rolling were linearlycorrelated with leaf water potentials. Resistance in paniclesincreased with progress decline in water potentials, but thechange was not as high as in leaves. Among test cultivars, diffusive resistance increased at thelate flowering stage but not at the early stage when panicleswere newly emerged. Increased water losses contributed to severespikelet desiccation and white heads. A few of the upland cultivarstested, such as Kinandang Patong, IRAT 140 and IRAT 13 conservedmore water and showed reduced water-deficit stress symptoms.Copyright1993, 1999 Academic Press Drought tolerance, drought susceptibility, rice panicles     

Effect of high temperature and water stress on pollen germination and spikelet fertility in rice

In future climates, rice could more frequently be subjected to simultaneous high temperature and water stress during sensitive developmental stages such as flowering. In this study, five rice genotypes were exposed to high temperature, water stress and combined high temperature and water stress during flowering to quantify their response through spikelet fertility. Microscopic analyses revealed significant differences in anther dehiscence between treatments and genotypes, with a moderately high association with the number of germinated pollen grains on the stigma. There was a strong relationship between spikelet fertility and the number of germinated pollen on stigmas. Although, all three stress treatments resulted in spikelet sterility, high-temperature stress caused the highest sterility in all five genotypes. A cumulative linear decline in spikelet fertility with increasing duration of independent high-temperature stress and in combination with water stress was quantified. Better anther dehiscence, higher in vivo pollen germination, and higher spikelet fertility were observed in both the N22 accessions compared with IR64, Apo and Moroberekan under high temperature, water stress and combined stress, indicating its ability to tolerate multiple abiotic stresses.     

Panicle Differentiation and Spikelet Number Related to Size of Panicle in Pennisetum americanum

The size of the developing panicle of pearl millet (Pennisetumamericanum (L.) Leeke) was studied during panicle differentiation(from panicle initiation to the completion of spikelet production)in plants grown in pots or in the field and supplied with varyinglevels of nitrogen. The duration of panicle differentiationrequired a constant thermal time (day degrees) under all nitrogensupplies. However, the rate of growth of the developing panicleduring this phase was retarded by low nitrogen supply. Duringpanicle differentiation, it appeared that the developing paniclehad to reach a critical size before developmental events suchas the initiation of spikelet primordia commenced; timing ofdevelopmental events was related to the size of the developingpanicle. The number of spikelets produced depended on the rate of growthof the differentiating panicle and the duration of the phaseof spikelet initiation (from appearance of the first spikeletprimordium to completion of spikelet differentiation). Low nitrogensupply reduced the number of spikelets produced, by retardingthe rate of growth of the differentiating panicle; this delayedthe time to initiation of spikelets and thereby reduced theduration of spikelet initiation. All spikelets (irrespectiveof nitrogen supply on the mainstem and on tillers) occupiedthe same area of panicle surface at the completion of differentiationof the panicle and at anthesis. Key words: Millet, Panicle differentiation, Spikelet number     

Sodium and potassium uptake of rice panicles as affected by salinity and season in relation to yield and yield components

Salinity is a major yield-reducing stress in many arid and/or coastal irrigation systems for rice. Past studies on salt stress have mainly addressed the vegetative growth stage of rice, and little is known on salt effects on the reproductive organs. Sodium and potassium uptake of panicles was studied for eight rice cultivars in field trials under irrigation with saline and fresh water in the hot dry season and the wet season 1994 at WARDA in Ndiaye, Senegal. Sodium and potassium content was determined at four different stages of panicle development and related to salt treatment effects on yield, yield components and panicle transpiration. Yield and yield components were strongly affected by salinity, the effects being stronger in the HDS than in the WS. The cultivars differed in the amount of salt taken up by the panicle. Tolerant cultivars had lower panicle sodium content at all panicle development stages than susceptible ones. Panicle potassium concentration decreased with panicle development under both treatments in all cultivars, but to a lesser extent in salt treated susceptible cultivars. Grain weight reduction in the early panicle development stages and spikelet sterility increase in the later PDS were highly correlated (p < 0.01) with an increase in panicle sodium concentration in both seasons, whereas reduction in spikelet number was not. The magnitude of salt-induced yield loss could not be explained with increases in sodium uptake to the panicle alone. It is argued that the amount of sodium taken up by the panicle may be determined by two different factors. One factor (before flowering) being the overall control mechanism of sodium uptake through root properties and the subsequent distribution of sodium in the vegetative plant, whereas the other (from flowering onwards) is probably linked to panicle transpiration. This revised version was published online in August 2006 with corrections to the Cover Date.     

Variable reproductive output among clones of Spartina alterniflora (Poaceae) invading San Francisco Bay,California: the influence of herbivory,pollination, and establishment site

Spartina alterniflora has recently been introduced to San Francisco Bay, California, and is rapidly invading open mud flats, growing in circular patches that we found to be individual genetic clones. We collected spikelet samples from more than 200 clones and observed germination rates ranging from 0% to 59%, indicating substantial variation in reproductive output among clones. Several experiments were performed to explore the cause of variation. Pollination manipulations showed that S. alterniflora is outcrossing, but pollen supplements did not increase spikelet germination rates. Exclusion of the only insect herbivore (a phloem feeder, Prokelisia marginata) from developing inflorescences increased the proportion of spikelets containing seed, but failed to increase germinations per spikelet. Spikelets from Willapa Bay, Washington, grown free of insect herbivores, had germination rates similar to San Francisco Bay. These results suggest herbivory is not limiting reproductive output of S. alterniflora. Spikelet viability was not related to clone size; however, clones located lower in the intertidal or far up a drainage slough averaged fewer germinations per spikelet, suggesting clones in areas with lower genet density may have lower spikelet viabilities. Spikelet samples from different sections of clones growing across wide environmental ranges had similar rates of germination, suggesting some genetic influence on spikelet viability. Differential reproductive output among clones and the novel selective environment of San Francisco Bay are expected to cause gene frequency changes in this rapidly expanding population.     

Brassinosteroids function in spikelet differentiation and degeneration in rice

Brassinosteroids(BRs) play crucial roles in many aspects of plant development. However, their function in spikelet differentiation and degeneration in rice(Oryza sativa L.) remains unclear. Here, we investigated the roles of these phytohormones in spikelet development in fieldgrown rice subjected to five different nitrogen(N)fertilization treatments during panicle differentiation. BR levels and expression of genes involved in BR biosynthesis and signal transduction were measured in spikelets. Pollen fertility and the number of differentiated spikelets were closely associated with 24-epicastasterone(24-epiCS) and28-homobrassinolide(28-homoBL) levels in spikelets.Enhanced BR biosynthesis and signal transduction, in response to N treatment, enhanced spikelet differentiation, reduced spikelet degeneration, and increased grain yield. Increases in proton-pumping ATPase activity, ATPconcentration, energy charge, and antioxidant system(AOS) levels were consistent with 24-epiCS and28-homoBL concentrations. Exogenous application of24-epiCS or 28-homoBL on young panicles induced a marked increase in endogenous 24-epiCS or 28-homoBL levels, energy charge, AOS levels, spikelet differentiation, and panicle weight. The opposite effects were observed following treatment with a BR biosynthesis inhibitor. Our findings indicate that, in rice, BRs mediate the effects of N fertilization on spikelet development and play a role in promoting spikelet development through increasing AOS levels and energy charge during panicle development.