92 resultados para Pollen grains
Resumo:
空中花粉是植被类型、气候状况与大气环境质量的指示物之一。空中花粉雨的取样实验对化石花粉谱的解释和古植被的恢复具有指导意义,因此空中花粉分析成为第四纪孢粉学和古生态学研究的有效辅助手段。本研究在新疆天山中段北坡进行了短期的空中花粉监测(2001年7月~2006年7月),目的是获得局域性和区域性植被所散发的花粉类型及其数量信息,并探讨花粉数量的植被与气候指示意义、花粉传播以及受环境条件影响的程度,同时分析和检测花粉浓度受气候条件的影响情况。空中花粉的收集使用Cour-type捕捉器,分别在3个不同海拔梯度进行5年连续的空中花粉收集(夏秋季节每周一个样品,冬春季节每两周一个样品)。取样地点分别是新疆天山中段北部的天池气象观测站(43°53'58.38"N, 88°07'15.75"E , 海拔1942.5m)、中科院阜康荒漠生态系统定位站(44°17'27.41"N, 87°55'52.65"E, 海拔477m)和准噶尔盆地古尔班通古特沙漠南缘的北沙窝试验站(44°22'40.74"N, 87°55'9.74"E, 海拔443m),取样点的植被带跨度天山北坡的森林草原到低海拔的典型荒漠。 光学显微镜鉴定的空中花粉主要有43种类型(24科30属),而蕨类、苔藓与真菌孢子数量较少。孢粉类型多样性指数为2.45(Shannon Index)。显然,从样品中所鉴定的空中花粉种类与所观察的植物种类有很大的差距。花粉类型与数量在三个不同海拔梯度的取样点上也有明显差异。天池取样点的花粉类型较丰富,与天山北坡垂直带植被分布以及丰富的植被成分相对应,花粉的优势种类为雪岭云杉(Picea schrenkiana)、蒿属(Artemisia)、藜科(Chenopodiaceae)、柽柳科(Tamaricaceae)和麻黄属(Ephedra),高代表性的外源荒漠花粉类型“削弱了”本地花粉类型的代表性;阜康与北沙窝取样点的花粉类型主要来自荒漠成分的藜科、蒿属、麻黄属、蒺藜科(Zygophyllaceae, 白刺属Nitraria为主)、柽柳科(柽柳属为主)、蓼科(Polygonaceae, 沙拐枣属Calligonum为主)等耐旱植物。花粉数量与植被数量并非简单的线性关系。空中和表土花粉的数量分析表明,藜科、蒿属、麻黄属花粉具有超代表性;雪岭云杉花粉的代表性适中;禾本科(Gramineae)、榆属(Ulmus)、白刺属、柽柳科、沙拐枣属花粉代表性较差。 5年平均花粉浓度大小依次是阜康(81.668 grains/100m3)>天池(51.726 grains/100m3)>北沙窝(45.7685 grains/100m3)。蒿属花粉的峰值出现在秋季,并且出现双峰现象(可能与不同蒿类花期分布不均有关),而其它木本和草本类植物花粉浓度的峰值为单峰,具有明显的季节性,春夏之交开花的桦木属、榆属等类型较为明显。藜科花粉的峰值出现在夏季,麻黄和柽柳的峰值出现在盛夏,桦木属、柳属Salix、雪岭云杉等花粉浓度峰值出现在初夏,与其花期物候相对应。木本植物花粉以雪岭云杉为主,其次为桦木属(Betula)、榆属、柳属、胡颓子科(Elaeagnaceae)花粉,柳属花粉百分比在三个取样点分配比较均匀,桦木属花粉在空间分布上差异明显,与植物分布数量相关。胡颓子科的花粉出现时间短,数量也较少。榆属花粉出现的季节性明显,代表性也较差。因此解释地层中出现类似的低代表性和高代表性的花粉类型时,需要注意其百分比数量的适当校正。木本植物花粉百分比随海拔高度变化成正比关系,对植被的指示性较好。雪岭云杉的年平均花粉浓度在天池取样点明显高于其它两个取样点,5年的平均花粉浓度和百分比与海拔高度呈正比。天池取样点的百分比最高为28.85%,平均为21.15%,而北沙窝与阜康两地5年平均百分比含量分别为0.69%和1.57%,这种时空变化规律与植被数量的关系密切。藜科花粉百分比在阜康和北沙窝取样点占绝对优势,5年的平均花粉浓度与海拔高度呈反比。蒿属花粉年平均浓度与海拔高度关系微弱(R2=0.04,p =0.46),而花粉百分比与海拔高度有显著的线性正比关系(R2=0.72,p<0.0001)。但在低海拔的两个取样点之间差异不明显。 年平均花粉百分比和花粉浓度随海拔梯度而变化,主要花粉类型的变化规律存在差异。暗针叶林雪岭云杉花粉的年平均百分比在低海拔的阜康和北沙窝地区低于3%,在盛花期的6月初,低海拔的取样点也未见超过5%,雪岭云杉花粉的传播在研究区范围内数量分布变化较大,再次表明该类型的花粉在原地沉积效率很高。空中花粉数量能够较好地指示主要的植被带,盛花期内50%以上的雪岭云杉花粉含量则指示了雪岭云杉森林植被带,而高含量的蒿属植物花粉指示了天山山前冲积平原上的蒿类荒漠,相反,高含量的藜科花粉代表了低海拔地区的荒漠。雪岭云杉花粉百分比与浓度均与海拔高度呈显著的线性正比关系,表明雪岭云杉花粉传播的局限性。主要的非木本植物藜科花粉的浓度与海拔高度成反比,蒿属则相反,表明藜科与蒿属花粉在研究区具有很好的植被与气候指示意义。 花粉绝对数量、百分比、浓度具有明显的年际变化,其原因与气候状况的变化有关,年平均花粉浓度、主要类型的花粉浓度与气候参数(降水量、平均温度、最高和最低温度、相对湿度等)的相关分析表明,降水量和湿度与花粉浓度呈负相关程度较高,与其它气候参数则呈显著的正相关。干旱指数、花粉比值、花粉百分比(AP,NAP)与七月份平均气温,年平均降水量的相关系数(Pearson correlation),Ar/Ch与七月平均温度在0.01水平上呈显著的正相关(R2=0.894), 与年平均降水量在0.01水平上呈显著的负相关(R2=0.874)。AP花粉百分比与七月份平均温度呈显著的负相关(R2=0.71, p<0.0001),与年降水量呈正相关(R2=0.43, p=0.01),而NAP花粉正相反。利用短期监测的空中花粉雨数据计算得到的花粉比值以及干旱指数在研究区具有明确的指示意义,尤其是干旱指数、Ar/Ch、Ar/Ep,这些都可为表土花粉、植被与气候关系模型的建立提供重要的参考信息。 气候参数在很大程度上影响大气花粉的浓度,而风速和风向对空中花粉的传播、散布影响较大,花粉的来源与传播受风向和风速的控制。不同取样点的空中花粉数量差异受地形条件影响很明显,这种差异也可利用地形空气动力学特征的差异来解释,天池监测站因“逆坡”(upslope)的气流促进了低海拔的花粉流向高海拔传播。 研究区的空中花粉数量信息是研究表土花粉、植被和气候关系以及第四纪古生态学的重要基础。本研究提高了对研究区空中孢粉的类型及其传播规律的认识和理解,增加了解空中花粉受环境影响的主要因素及其影响程度,对地层孢粉类型鉴定和花粉数据的解释提出指导性建议,对本地区的古植被与古环境研究增加现代孢粉学的参考依据和信息,并可为建立区域性的现代孢粉-植被-气候关系模型提供可靠依据。 同时,本报告也讨论了存在问题和不足,并提出了今后工作的可能完善方案。
Resumo:
Two strong luminescence bands were observed from a-SiOx:H in the spectral range of 550-900 nm at room temperature. One is a main broad peak which blueshifts with oxygen content and the other is a shoulder fixed at about 835 nm. In conjunction with TR and micro-Raman spectra, we have proposed that the main band may originate from the amorphous silicon grains embedded in SiOx network, while the shoulder might be due to some defects induced by excess-silicon in these films. (C) 1997 Elsevier Science Ltd.
Resumo:
In this work, the formation and characterization of nano-sized grains on the modified surfaces of GCr15 and H13 steels have been investigated. The material was processed by pulsed laser surface melting (LSM) under different depths of de-ionized water. The microstructures and phases of the melted zones were examined by x-ray diffraction, environmental field emission scanning electron microscopy and high resolution transmission electron microscopy. The results indicate that LSM under water can successfully fabricate nano-scaled grains on the surfaces of steel, due to the rapid solidification and crystallization by heterogeneous nucleation. The elemental segregation of chromium and activated heterogeneous nucleation mechanism of austenite in liquid metal play a key role in the formation of nano-sized grains at high cooling rates. This one-step technique provides us a new way to prepare uniform nano-scaled grains, and is of great importance for further understanding the growth of nano-materials under extreme conditions.
Resumo:
瑞香狼毒(Stellera chamaejasme L.)是瑞香科(Thymelaeaceae)狼毒属的一种多年生野草,有毒。据调查,从20 世纪60 年代开始至今,狼毒在青藏高原东缘的高寒草甸上不断蔓延、密度不断变大,在一些地段甚至成为优势物种。有关狼毒在高寒草甸蔓延的生态系统效应的研究尚未见报道。本文从系统碳、氮循环的角度,分别研究狼毒在生长和非生长季节对高寒草甸生态系统的影响。同时,从花粉化感的角度,深入研究狼毒对当地同花期物种有性繁殖的影响。系统地研究高寒草甸生态系统物质循环过程,特别是非生长季节微生物和土壤碳氮库的动态变化,有助于揭示狼毒在系统物质循环方面的“物种效应”以及这种效应的季节变化,为丰富有关高海拔生态系统,特别是其非生长季的物质循环的科学理论做出贡献。同时,碳氮循环和花粉化感的研究还有助于深刻地理解狼毒作为一种入侵性很强的杂草的特殊的蔓延机制,从而为狼毒的有效防治、高寒草甸的科学管理提供依据。 针对狼毒在青藏高原高寒草甸上蔓延的生态系统碳氮循环方面的影响,开展以下2 方面的研究:(1)在生长季,研究松潘县尕米寺附近(北纬32°53',东经103°40',海拔3190 m)的两种地形(平地和阳坡)条件下狼毒对土壤碳氮循环影响及可能的原因。狼毒和其它几个主要物种(圆穗蓼(Polygonummacrophyllum D. Don var. Macrophyllum),草地早熟禾(Poa pretensis L.),四川嵩草(Kobresia setchwanensis Hand.-Maizz.),鹅绒委陵菜(Potentilla anserina L.var. anserine)和鳞叶龙胆(Gentiana squarrosa Ledeb.)的地上凋落物产量以及地上凋落物和根的化学组成被测量。在有-无狼毒斑块下,各种土壤的库(比如,铵态氮、硝态氮、无机磷和微生物生物量)和周转率(包括净矿化、净硝化、总硝化、反硝化和微生物呼吸速率)被测量和比较。(2)在非生长季节,尤其是春季冻融交替期,选取了两个研究地点——尕米寺和卡卡沟(北纬32°59',东经103°41',海拔3400 m),分别测定有狼毒和无狼毒斑块下土壤微生物生物量碳和氮、可溶性有机碳和氮以及铵态氮和硝态氮的动态变化。同时,分别在上述两个地点有-无狼毒的样地上,研究6 个主要物种(狼毒、圆穗蓼、草地早熟禾、四川嵩草、鹅绒委陵菜和鳞叶龙胆)从秋季开始、为期1 年的凋落物分解过程。 针对狼毒花粉化感对同花期其它物种可能的花粉化感作用开展以下工作:在实验室中,用一系列浓度的狼毒花粉水浸提液对与它同花期的其它物种以及自身花粉进行测试,测定花粉萌发率;在野外自然条件下的其它物种的柱头上施用上述浓度的狼毒花粉水浸提液,观测种子结实率,同时,观察狼毒花粉的种间花粉散布数量。 生长季节的研究结果表明,狼毒地上凋落物氮含量比其它几个主要物种更高,而木质素-总氮比更低。狼毒显著地增加其斑块下表层土壤中有机质的含量,而有-无狼毒的亚表层土壤在有机碳和总磷方面没有显著差异。狼毒表土中硝态氮含量在平地和阳坡比无狼毒土壤分别高113%和90%。狼毒表土中微生物生物量碳和氮量显著高于无狼毒表土。无论是平地还是阳坡,狼毒土壤的总硝化和微生物呼吸速率显著高于无狼毒土壤;而它们的反硝化速率只在平地有显著的差异。狼毒与其它物种间地上凋落物的产量和质量的差异可能是导致有-无狼毒土壤碳氮循环差异的原因。我们假设,狼毒可能通过增加贫氮生态系统土壤中的有效氮含量提高其入侵能力。 非生长季的研究结果表明,在青藏高原东缘的高寒草甸上,土壤微生物生3物量在11 月的秋-冬过渡期达到第一个峰值;在春季的冻融交替期,微生物生物量达到第二个峰值后又迅速降低。无机氮以及可溶性有机碳氮与微生物生物量有相似的变化过程。微生物碳氮比呈现显著的季节性变化。隆冬季节的微生物生物量碳氮比显著高于生长旺季的微生物碳氮比。这种变化可能暗示冬、夏季微生物的群落组成和对资源的利用有所不同。有-无狼毒斑块下土壤微生物和土壤碳、氮库一般只在秋-冬过渡期有显著差异,有狼毒土壤微生物生物量和土壤碳、氮库显著高于无狼毒土壤;而在之后的冬季和春季没有显著差异。所有6 个物种凋落物在非生长季分解率为24%-50%,均高于生长季的10%到30%。其中在秋-冬过渡期,凋落物开始埋藏的两周时间内,分解最快,达10%-20%。不同物种凋落物全年的分解率和分解过程有显著差异。圆穗蓼在全年的分解都较缓慢(非生长季26%,生长季15%),草地早熟禾和四川嵩草等全年的分解速率比较均匀(非生长季和生长季均为30%,非生长季略高),而狼毒在非生长季分解较快(约50%),而在接下来的生长季分解变得缓慢(约12%)。所有物种的凋落物氮含量在非生长季下降,而在随后的生长季上升。 实验室的花粉萌发试验证明,狼毒花粉对自身花粉萌发没有自毒作用,而其它受试的所有物种(圆穗蓼,秦艽(Gentiana macrophylla Pall. var. fetissowii),湿生扁蕾(Gentianopsis paludosa (Hook. f.) Ma var. paludosa),鳞叶龙胆,椭圆叶花锚(Halenia elliptica D. Don var. elliptica),蓝钟花(Cyananthus hookeri C. B.Cl. var. grandiflorus Marq.),小米草(Euphrasia pectinata Ten.),川西翠雀花(Delphinium tongolense Franch.),高原毛茛(Ranunculus tanguticus (Maxim.)Ovcz. var. tanguticus)和鹅绒委陵菜)的花粉萌发率随着狼毒花粉浸提液浓度的增加呈显著的非线性降低。大约3 个狼毒花粉的浸提液就可以抑制受试的多数物种的50%的花粉萌发。在鳞叶龙胆和小米草柱头上狼毒花粉的数量分别为5.76 个和3.35 个。狼毒花粉散布数量的差异最可能的原因在于是否有共同的传粉昆虫。花的形状(辐射对称VS 左右对称)、植株或花的密度以及花期重叠性可以部分解释这种差异。在野外试验中,我们发现6 个物种(秦艽、湿生扁蕾、鳞叶龙胆、椭圆叶花锚、蓝钟花和小米草)的种子结实率随狼毒花粉浸提液浓度的增加呈显著的非线性降低。鳞叶龙胆和小米草柱头上狼毒花粉的数量(分别是5.76 个和3.35 个)分别达到了抑制它们63%和55%种子结实率的水平。因此,狼毒对鳞叶龙胆和小米草可能存在明显的花粉化感抑制作用。狼毒周围的物种可能通过花期在季节或昼夜上的分异避免受到狼毒花粉化感的影响或者通过无性繁殖来维持种群繁衍,因此狼毒通过花粉化感作用对其周围物种繁殖的影响程度还需要进一步地研究。如果狼毒的花粉化感抑制作用确实存在,那么它可能成为一种自然选择压力,进而影响物种的进化。 Stellera chamaejasme L., a perennial toxic weed, has emerged and quicklydominated and spread in the high-frigid meadow on the eastern Tibetan Plateau ofChina since the 1960s. In the present study, effects of S. chamaejasme on carbon andnitrogen cycles on the high-frigid meadow on the eastern Qinghai-Tibetan Plateau ingrowing and non-growing season, and its pollen allelopathic effects on the sympatricspecies were determined. The present study that focused on carbon and nitrogencycles, especially on microbial biomass and pools of carbon and nitrogen innon-growing season, could profoundly illuminate plant-species effects on carbon andnutrient cycles and its seasonal pattern and help to understand spread mechanism ofS. chamaejasme as an aggressive weed. The present study also contributed to furtherunderstand carbon and nutrient cycles on alpine regions in non-growing season andprovide a basis on weed control of S. chamaejasme and scientific management in thehigh-frigid ecosystem. Effects of S. chamaejasme on carbon and nitrogen cycles on the high-frigidmeadow on the eastern Qinghai-Tibetan Plateau were determined. The study couldbe divided into two parts. (1) In the growing season, we quantified the effects of S.chamaejasme on carbon and nitrogen cycles in two types of topographic habitats, theflat valley and the south-facing slope, where S. chamaejasme was favored to spreadlitter and root were measured to explain the likely effects of S. chamaejasme on soilcarbon and nutrient cycles. The sizes of various soil pools, e.g. nitrite, ammonium,inorganic phosphorus and microbial biomass, and turnover rates including netmineralization, gross nitrification, denitrification and microbial respiration weredetermined. (2) In the non-growing season study, microbial biomass carbon andnitrogen, soluble organic carbon and nitrogen, ammonium and nitrate weredetermined through the non-growing season, especially in the processes offreeze-thaw of spring in two high-frigid sites, i.e. Kaka valley and Gami temple, onthe eastern Qinghai-Tibetan Plateau. Meanwhile, litter decomposition of six commonspecies, including Stellera chamaejasme L., Polygonum macrophyllum D. Don var.Macrophyllum, Poa pretensis L., Kobresia setchwanensis L., Potentilla anserina L.var. anserine and Gentiana squarrosa Ledeb., were also examined under theabove-mentioned experimental design through one whole-year, which began in theautumn in 2006. In the study of pollen allelopathy, several work, including in vitro study oneffects of extract of pollen from S. chamaejasme on sympatric species and pollenfrom itself, field experiments on effects of pollen extract with the same regime ofconcentrations on seed set and field observation on heterospecific pollen transfer ofS. chamaejasme to six of those sympatric species has been done. The results in the growing season showed that aboveground litter of S.chamaejasme had higher tissue nitrogen and lower lignin: nitrogen ratio than thoseco-occurring species. S. chamaejasme significantly increased topsoil organic matter,whereas no significant differences were found for organic C and total P in subsoilbetween under-Stellera and away-Stellera locations. The nitrate in Stellera topsoilwas 113% and 90% higher on the flat valley and on the south-facing slope,respectively. Both microbial biomass C and N were significantly higher in Stelleratopsoil. Gross nitrification and microbial respiration were significantly higher inStellera topsoil both on the flat valley and on the south-facing slope, whereassignificant differences of denitrification were found only on the flat valley. Thedifferences in the quantity and quality of aboveground litter are a likely mechanismresponsible for the changes of soil variables. We assumed that S. chamaejasme couldenhance their spread by increasing nutrient availability in N-deficient ecosystems. The results in the non-growing season showed that microbial biomass achievedthe first summit in late autumn and early winter on the eastern Qinghai-TibetanPlateau. In the stages of freeze-thaw of spring, microbial biomass firstly achieved thesecond summit and subsequently sharply decreased. Inorganic nitrogen, solubleorganic carbon and nitrogen had a similar dynamics with that of microbial biomass.Ratio of microbial biomass carbon and nitrogen had an obviously seasonal pattern.The highest microbial C: N were in the non-growing season, which weresignificantly higher than those in the growing season. The seasonal pattern inmicrobial biomass C: N suggested that large changes in composition of microbialpopulation and in resources those used by microbes between summer and winter.Generally, microbial biomass and pools size of carbon and nitrogen in Stellera soilwere significantly higher than those under adjacent locations in late autumn andearly winter, but there were not significant differences in winter and in spring. Litterof all the focal species (Stellera chamaejasme L., Polygonum macrophyllum D. Donvar. Macrophyllum, Poa pretensis L., Kobresia setchwanensis Hand.-Maizz.,Potentilla anserina L. var. anserine and G. squarrosa Ledeb.) decomposed about24%-50% in the non-growing season, which were higher than those in the growingseason (ranged from 10% to 30%). Litter decomposed 10%-20% within the first twoweeks in late autumn and early winter. Significant differences in the whole-yeardecomposition rate and in the processes of decomposition were found among species.Polygonum macrophyllum decomposed slowly through the whole year (26% and15% in the non-growing season and in the growing season, respectively). Certainspecies, such as P. pretensis L. and K. setchwanensis, decomposed at a similar rate(30% both in the non-growing and in the growing season, slightly higher in the8growing season than those in the growing season), whereas S. chamaejasmedecomposed more rapidly (about 50%) in the non-growing season and subsequentlydecomposition became slow (about 12%) in the growing season. Litter nitrogencontents of all the focal species firstly decreased in the non-growing season and thenincreased in the growing season. In vitro experiments of pollen allelopathy, the results showed that pollen from S.chamaejasme was not autotoxic, whereas pollen germination in all the sympatricspecies (Polygonum macrophyllum D. Don var. Macrophyllum, Gentianamacrophylla Pall. var. fetissowii, Gentianopsis paludosa (Hook. f.) Ma var. paludosa,Gentiana squarrosa Ledeb., Halenia elliptica D. Don var. elliptica, Cyananthushookeri C. B. Cl. var. grandiflorus Marq., Euphrasia pectinata Ten., Delphiniumtongolense Franch., Ranunculus tanguticus (Maxim.) Ovcz. var. tanguticus andPotentilla anserina L. var. anserina) decreased nonlinearly as the increasingconcentrations of extract of pollen from S. chamaejasme. Pollen Extract of threepollens from S. chamaejasme generally inhibited 50% pollen germination of most ofthe focal species. 5.76 and 3.35 pollens from S. chamaejasme were observed in fieldon stigmas of G. squarrosa and E. pectinata, respectively. Differences inheterospecific pollen transfer of S. chamaejasme could be attributed to the primaryreason whether they shared common pollinators. Flower morphology (e.g.zygomorphic or actinomorphic), plant or floral density and concurrence in floweringphonologies could explain, in part, the differences in heterospecific pollen transfer.In field experiments, the results showed that seed set in six sympatric species(Gentiana macrophylla Pall. var. fetissowii, Gentianopsis paludosa (Hook. f.) Mavar. paludosa, Gentiana squarrosa Ledeb., Halenia elliptica D. Don var. elliptica,Cyananthus hookeri C. B. Cl. var. grandiflorus Marq. and Euphrasia pectinata Ten.)decreased nonlinearly as the increasing concentrations of extract of pollen from S.chamaejasme. According to the nonlinear curves, the amounts of pollens from S.chamaejasme on stigmas of G. squarrosa and of E. pectinata (i.e. 5.76 grains and3.35 grains, respectively) could reduce 63% and 55% seed set of G. squarrosa and ofE. pectinata, respectively. Thus, allelopathic effects of S. chamaejasme on G.squarrosa and E. pectinata could be realistic. The sympatric species of S.chamaejasme could avoid pollen allelopathy of S. chamaejasme to sustainthemselves. This highlights the need to study how much pollen allelopathy in S.chamaejasme influences the sympatric species through divergence in seasonal ordiurnal flowering phonologies or through asexual reproduction. If pollen allelopathyin S. chamaejasme was confirmed, it could be as a pressure of natural selection andthus play an important role in species evolution.
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水稻是重要的粮食作物,其产量的增加和品质的改良都是关系国计民生的大事。就我国现阶段的国情而言,水稻产量在现有水平上稳步提升仍是未来十几年甚至几十年农业生产最重要的目标之一。尽管根据“超级杂交水稻育种”的战略设想和水稻育种实践,通过不断地改进育种技术可望在更高的产量水平上进行水稻杂种优势利用,在稻属植物内还具有很大的产量潜力可以挖掘。然而,仅仅从现有的种质基础出发,要更大幅度提高水稻单产,实现“超级杂交稻”的目标也存在一些困难:现有的推广品种是二倍体,尽管种类众多,但是其基因组的来源相对单一;同时,水稻基因组DNA含量也是作物中最少的,基因组内寻求开发潜力有一定困难;水稻作为C3植物,光合利用效率不高也是制约水稻产量提高的因素之一。因此,寻求常规手段以外的技术突破或者方法创新,是实现“超级杂交稻”的目标的迫切需求。本研究利用秋水仙素能抑制细胞分裂中纺锤丝的收缩、使细胞染色体加倍的作用,对水稻幼穗诱导的愈伤组织细胞进行加倍,并分化出再生植株;创制出水稻同源四倍体新的种质材料,在此基础上选育水稻同源四倍体雄性不育三系材料,并实现水稻同源四倍体的三系配套,开展水稻同源四倍体杂种优势利用和四倍体杂交水稻选育研究,建立水稻同源四倍体杂种优势利用的新技术体系。这不仅有助于倍性水平杂种优势的开拓和利用,同时也将为我国新世纪“超级稻”育种研究开辟一条新的技术途径。 水稻幼穗诱导愈伤组织并分化成苗是一项成熟、简单的组织培养技术。本研究以普通二倍体水稻亲本为材料,用秋水仙素进行水稻的多倍体化诱导,创制同源四倍体水稻三系亲本材料并对其进行鉴定。多倍体化以秋水仙素诱导的愈伤组织培养为基础,研究不同秋水仙素浓度梯度和愈伤组织诱导培养基组合对诱导四倍体植株的影响。结果表明在MS+2,4 D 1.0mg/L+ KT0.2mg/L+ IAA0.2mg/L 和500mg/L的秋水仙素处理下,水稻愈伤组织染色体加倍(有最高的效率)效果较好,平均加倍频率可达25.26%,其中,材料CDR22和IR26诱导较易成功,加倍频率分别达到75%和26.5%;相对材料94109 1.3%加倍频率和冈46B 10.8%加倍频率,诱导率差异极显著。 对水稻四倍体材料进行了形态学鉴定结果表明,与二倍体水稻对照相比其株高、穗长、花粉育性等主要农艺性状,确定四倍体材料在穗长和千粒重两方面极显著提高,种子的长度和宽度也显著增长。对花粉育性鉴定,确认水稻四倍体不育系材料仍为不育,保持系材料自交和杂交可育,恢复系材料自交和杂交可育。对四倍体材料进行细胞形态、染色体数目等方面进行细胞学鉴定,经核型分析表明水稻四倍体材料具有48条染色体,是二倍体水稻的两倍。水稻四倍体材料根尖分生组织细胞与二倍体的根尖分生组织细胞相比,细胞体积、细胞核和核仁显著增大。四倍体三系材料在细胞有丝分裂中期均可规则排列在赤道板,并能均等地移向两极;后期观察中没有发现染色体分离滞后现象,分裂末期细胞能够形成大小相对均一的子细胞。水稻同源四倍体三系材料细胞分裂未见异常,植株生长发育正常。 从1996年至2006年,针对结实率、有效分蘖、着粒数和穗长等主要农艺性状,通过系谱选育的方法,对培育的同源四倍体水稻亲本材料进行了连续选择和改良,取得较好成效。表现为结实率的改良效果极佳,所有改良材料的平均结实率均呈上升趋势,如D237(29.70%→72.70%)、DTB(19.55%→53.21%)等。有效分蘖总体呈现上升趋势,但在不同的年份,如1998和2002存在较大的负向波动。部分材料改良效果明显,如D19B(5.87→13.50)、D什香 (7.00→12.00)等;同时一些材料如DTB和D明恢63虽然总体略有提高,但在不同的年份波动很大,因此存在较大改良阻力,原因还有待进一步研究。着粒数的改良上升趋势比较显著,除保持系的DTB之外,其余材料的平均着粒数有显著提高。穗长的改良阻力较大,虽然不同材料总体上有所提高,但效果并不显著,并且不同年份有较大负向波动(2001)。此外还对株高、剑叶长等性状也进行了选择,但效果不显著,原因有待进一步提高。同源四倍体材料产量相关性状遗传改良幅度不一致,保持系和恢复系间的遗传改良效果也存在差异。这为同源四倍体水稻的进一步利用打下了良好的基础。 籼稻和粳稻亚种间杂交及杂种优势利用的主要障碍就是其低的结实率。而同源四倍体杂交水稻的研究为提高杂交水稻的杂种优势利用创造了新的途径。本研究通过随机区组设计方案,挑选性状优良的二倍体水稻材料,包括雄性不育系,保持系和恢复系进行秋水仙素诱导加倍,从而获得同源四倍体水稻对应的三系材料。利用选育的优良水稻同源四倍体三系材料,配制7个杂交组合,杂交F1代与其恢复系亲本进行比较,用于计算超亲优势(HB);而杂交F1代与生产上大面积推广的二倍体杂交品种汕优63进行比较,用于计算杂种优势。结果显示,同源四倍体杂交水稻的超亲优势表现为:每株有效穗变化幅度为1.4%至105.9%,总粒数为0.5%至74.3%,每穗实粒数为17.6%至255.7%,结实率为9.6%至130.4%。这些农艺性状的改良使得这7个杂种F1的理论产量的超亲优势高达64.8%至672.7%。小区试验中四倍体杂交水稻组合T461A/T4002和T461A/T4193分别比二倍体对照汕优63提高46.3%和38.3%以上,除一个品种以外所有品种产量均接近或高于汕优63的产量。同源四倍体水稻强大的杂种优势表明,亚种间杂交育性低的问题可通过四倍体化及强化选择来解决。此外,同源四倍体杂交水稻器官的巨大性也是其产量提高的有利因素,水稻同源四倍体三系杂种优势利用研究具有一定的理论价值和商业生产潜力。 Rice is one of the major food crops, the improvement of the production and quality of it is an important thing related to the people's livelihood. On China's current national conditions, steadily increase of the rice yield based on the current level is still one of the most important goals in the next decade or even decades of agricultural production. According to the "super hybrid rice breeding" the strategic and rice breeding practice, improvement of the use of hybrid rice heterosis through continuous improvements in breeding technology is expected to get a higher level of rice yield, there are also a great yield potential can be exploited. However, there are also some difficulties to increase rice yield obviously and implement the goal of "super hybrid rice" based on the existing germplasm: Rice varieties in promotion are diploid, although there are many varieties, but their genome are from a comparatively single source; Meanwhile, the rice genome DNA are the least among the crops, it is difficult to exploit the development potential within the genome; Rice as C3 plants, photosynthetic efficiency is not high, it is one of the factors constraint rice yield. Therefore, seeking technological breakthroughs or innovative methods different from conventional means is the urgent needs to reach the target of "super hybrid rice". Using colchicine inhibit spindle contraction during cell division, double the cell chromosome, we induced callus cells from rice panicle to be doubled, and differentiated regeneration; we created a new autotetraploid rice germplasm material, and on that basis we bred male sterility three line autotetraploid rice materials, and the achieved the three line rice autotetraploid matchmaking, researched in autotetraploid rice heterosis usage and tetraploid hybrid rice breeding, constituted a new technology system of autotetraploid hybrid rice heterosis utilization. This not only helps the tetraploid rice heterosis exploration and use, but also inaugurates a new technical means for China in the new century "super rice" breeding research. We chose ordinary diploid rice as materials, using colchicine to induce the polyploidization, created the autotetraploid rice three-line materials and identified them. The polyploidization was based on the colchicine-induced callus tissue culture, and we experimented different colchicine concentrations and culture mediums to induce tetraploid plants, confirmed that the optimal concentration for inducement was 500 mg/L, the average induce rate was 25.26 %. Among all the materials, CDR22 and IR26 had higher induced rate; in contrary, 94109 and GANG46B had lower induced rate, the difference was significant. Autotetraploid materials was identified of both morphological and cytological, compared plant height, length of pollen sterility, and other major agronomic traits with a diploid rice as the control plant, identified that the autotetraploid materials had very significant advantages in ear length and thousand-grain weight, as well as the size of the seeds. Cytology identification included observation of the cell morphology, the number of chromosomes, and karyotype analysis on the autotetraploid materials confirmed that their chromosome number was 48, twice of the diploid rice. Mitoses in the three lines were common: chromosomes arrayed normally in metaphase and separated balanced into the two poles, chromosome moved without lagging in anaphase and daughter cells normally formed in telophase except one. It has been proved that tetraploid rice has normal meiosis as their diploid relatives, which usually including series of sub-phases as interphase, prophase I (five sub-phases), prophase II, metaphase I, II, anaphase I, II and telophase I, II. However, abnormal phenomena, such as formation of tetravalent, trivalent and univalent, chromosome lagging and so on, which would finally block meiosis. Configurations of chromosome in metaphaseⅠwere versatile in structure and form accept the bivalent. That condition varied in different strain, suggesting more complex paring configurations and more versatile genetic characters in tetraploid rice. All these abnormalities in meiosis contributed to low fertility of gamete and might consequently resulted in low seed setting. Successive selection and improvement on seed set, productive tiller per plant, total grains per panicle, panicle length and so on had been carried out from 1996 to 2006. The raise of seed sets was significant in both restorers and maintainers. Seed sets of some strains were improved more significantly than others, for example D237(29.70%→72.70%)、DTB(19.55%→53.21%)and et al.. Productive tiller per plant was improved to some extant. The tendency of improvement was rising on the whole but changed in some years such as 1998 and 2002. Part of the stains increased greatly, such as D19B(5.87→13.50)、Dshixiang (7.00→12.00) and so on, but some strains including DTB and Dminghui63 only increased little and decreased in some years by unknown reason. Total grains per panicle increased significantly and all strains except DTB increased. Improvement of panicle length termed to be hard. Different strains showed different capacities for improvement and floating existed in different years for example 2001. It has been proved that other agronomical traits including plant length, flag leaf length and so on could be improved but not significantly by selection also. In a word, agronomical traits could be raised by successive selection that is prerequisite for further utility of autotetraploid rice. Poor fertility is the main barrier for utilizing heterosis between the two rice (Oryza stiva L.) sub-species, indica and japonica. Recently, the development of autotetraploid hybrids (2n=4x=48) has been suggested as a new method for increasing heterosis in hybrid rice. Using standard experimental protocols, the elite diploid rice male sterile, maintainer, and restorer lines were colchine-doubled and autotetraploid counterparts were obtained. Seven resulting hybrids were analyzed for heterobeltiosis (HB), where the F1 was compared to the male parent, and the degree of heterosis, where the F1 was compared to the diploid commercial hybrid, Shanyou 63. The HB among the autotetraploid hybrids ranged from 1.4 to 105.9% for the productive panicles per plant, 0.5 to 74.3% for total kernels per panicle, 17.6 to 255.7% for filled kernels per panicle, and 9.6 to 130.4% for seed set. Improvements in these yield components resulted in the HB for kernel yield ranging from 64.8 to 672.7% among the seven hybrids. Hybrids T461A/T4002 and T461A/T4193 yielded 46.3 and 38.3% more, respectively than Shanyou 63, and all other hybrids but one yielded the same or more than Shanyou 63. The high heterosis for yield suggests that hybrid sterility between two rice sub-species may be overcome by using tetraploid lines followed by intensive selection. Also, the gigantic features of the autotetraploid hybrids may establish a plant structure able to support the higher yield.
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课题组在不断地创制新的同源四倍体材料的同时,连续多年以提高结实率为目的培育、筛选自交系材料,已获得自交繁殖十二年的高代自交系材料。相对于诱导创制初期,材料表现出的结实率低,同种系单株间的差异较大;高代材料已表现出较显著的结实率提升和较一致的农艺性状表型。 本实验选取课题组多年培育的同源四倍体水稻高代自交系材料,通过形态学、农艺性状和细胞遗传学比较,研究水稻同源四倍体与二倍体之间的异同。结果显示,所有同源四倍体材料的染色体组成均为2N=4X=48,花粉母细胞(PMC)减数分裂行为较正常,99%以上的染色体都能在减数分裂中期I(MI)发生联会、配对,形成四价体和二价体,这与理论染色体组构成相符。在减数分裂过程中,结实率较高的材料染色体异常现象较少而结实率较低的材料染色体异常现象较严重。统计分析表明,二价体和四价体的比例对结实率没有显著影响,但是三价体的数目对结实率有一定影响。这一结果表明了结实率和细胞减数分裂行为可能存在相关性,同源四倍体的减数分裂行为为筛选高结实率的同源四倍体种系提供了依据。 然后,对同源四倍体水稻高代自交系材料进行农艺性状和品质性状的统计与分析。主要针对结实率、每穗实粒数、有效分蘖和穗长等主要农艺性状,以及直链淀粉含量这一重要的品质性状进行统计。将统计结果与1996年诱导加倍的初代材料的数据相对比分析,结果显示所有材料经过多代选育培养,其农艺性状已经有了较显著的提高,同时同源四倍体材料的农艺性状稳定性也有了较显著的提升。如结实率的提高幅度较大,所有材料的平均结实率均显著高于加倍初代,而同种材料不同单株间的结实率差异也显著地减少,变异系数(CV)的平均值由1996年的41.15%减少到了2008年的28.81%。其他重要农艺性状也有不同程度的改良,种内变异系数也相应地降低。此外,实验测量了同源四倍体材料和来源二倍体材料的直链淀粉含量。分析结果显示,部分材料的直链淀粉含量与二倍体亲本产生了较显著的差异,这可能是诱导加倍过程中的遗传变异造成的;同源四倍体材料的种内变异系数(CV)平均值由1996年的6%下降到了2008年的3.88%,显示出在品质性状方面,同源四倍体材料的遗传稳定性也有较显著的增加。同源四倍体材料农艺性状经过多年的选育,表现出一定的提升,同时,经过多年自交纯化,所有材料种系内的性状差异逐渐缩小,说明同源四倍体水稻的遗传稳定性随着自交纯化而增强,这为同源四倍体水稻的进一步选育打下了良好的基础。 最后,通过测量连续两年的自交系材料的遗传多态性,分析材料间遗传差异和种群遗传结构,深入研究连续两代材料间的遗传差异,研究同源四倍体水稻与二倍体材料遗传稳定性之间的差异。实验采用18对SSR微卫星标记对连续两代15个材料,共94份样本进行差异分析。通过扩增条带长度多态性分析,计算不同材料以及同种材料不同世代间的遗传距离,构建同源四倍体和二倍体水稻的分子指纹库,并绘制聚类图。结果显示,同源四倍体和二倍体不同材料间的遗传差异比较大,遗传距离处于0.4757至0.2816之间;而相同品种不同世代材料间的遗传差异较小,但也表现出一定的遗传差异。同种同源四倍体材料不同世代间的遗传差异比二倍体材料更大,两代四倍体材料间遗传距离处于0.1359至0.0485之间;而两代二倍体材料间的遗传距离处于均小于0.0388。结果表明,同源四倍体水稻高代材料具有一定的遗传稳定性,但与来源二倍体材料相比,其世代间的遗传变异性仍然较强。这种结果说明,经过多代的自交纯化培育,同源四倍体水稻材料能够建立起相对稳定的遗传结构,同时,其强于二倍体亲本的变异性有能够为新品种的选育,农艺性状、品质性状的改良提供一定的遗传基础。此外,分析结果表明通过分子标记辅助检验,水稻材料间的遗传多态性能够有效地区分不同的品种,这为水稻品种的分子鉴定提供了一定的依据。 本研究从细胞学鉴定,农艺性状统计分析以及分子标记辅助聚类分析多方面地对同源四倍体水稻高代系进行了研究,对探究同源四倍体水稻的遗传规律,进一步揭示其遗传特性、农艺性状的遗传构成,为进一步选育优质的多倍体水稻提供了一定的理论依据。 This group insists on creating new Autotetraploid Rice (Oryza sativa L.) materials, while improving the seed-setting of them for many years, cultivated and selected the inbred line materials, has obtained the high generation inbred lines after twelve years cultivation. Compared to the early induced materials, which shown the low seed setting, and the large difference between the different plants in the same germ-line; the high generation materials have shown significant improvement in seed setting and more uniform phenotype agronomic traits. The autotetraploid rice high generation inbred lines material, which has been cultivated for more than 12 years, was chose in this experiment. The similarities and differences between autotetraploid and diploid rice was studied through morphological, agronomic and cytogenetic ways. The results showed that all the chromosome of autotetraploid materials are composed of 2N=4X=48, the pollen mother cells (PMC) meiosis behavior is normal, more than 99% chromosomes in metaphase I(MI) were federated and paired to form tetravalents or bivalents, which constitutes a consistent theory of genome. In the meiosis process, the material with a higher seed setting showed less chromosome abnormal than the material whose seed setting is lower. However, statistical analysis showed that the bivalent and tetravalent rate had no significant impact on seed setting, but the number of trivalent had a certain impact on seed setting. The result shows that the seed setting may be related to the meiosis behavior, which provides a basis to cultivate new autotetraploid germ line with high seed setting through the meiotic behavior. Furthermore, the agronomic and quality traits of autotetraploid rice high generation inbred material were statistically analyzed. The statistically analysis was focused on major agronomic traits such as: seed setting, grains per panicle, effective tillers and panicle length, as well as the important quality trait amylose content. The statistic data was compared with the data in 1996, when the first induced generation of autotetraploid material, and the result shows that after a multi-generation breeding, the agronomic traits has been significantly improved in all the materials, while the stability of agronomic traits also significant upgraded. For instant, the seed setting increased significantly, the average seed setting of all materials was significantly higher than the first induced generation, and the differences between different plants in the same species also significantly reduced, the average of the coefficient of variation (CV) was reduced from 41.15% in 1996 to 28.81% in 2008. Other important agronomic traits had improved in different degrees; the coefficient of variation within species is also reduced accordingly. In addition, the amylose content of autotetraploid and diploid materials was measured in this experiment. The results shows that the amylose content of some of the material differed from diploid parents significantly, it may caused by the genetic change during the inducing, autotetraploid materials intra-specific coefficient of variation (CV) average reduced from 6% in 1996 to 3.88% in 2008, shows that this is a significant increase of quality traits stability in autotetraploid rice. Agronomic traits of autotetraploid material shows some improvement after years of breeding, at the same time, after years of purification, all material within the germ-line gradually narrow the differences in traits indicates that autotetraploid rice genetic stability was enhanced, which laid a good foundation for the further autotetraploid rice breeding. Finally, this experiment studied the genetic differences between materials of two generations and researched the difference of genetic stability between diploid and autotetraploid rice materials through investigating the genetic polymorphism, genetic differences between materials and population genetic structure of inbred line materials of two consecutive years.18 pairs of SSR microsatellite markers for 15 materials of two generations were used in this experiment, and the total of 94 samples were analyzed. Through the amplification length polymorphism analysis of different materials and materials in different generations, the genetic distance between materials and generations was analyzed, a diploid and autotetraploid rice molecular fingerprint database and map rendering cluster were constructed. The result shows that the genetic distance is between 0.4757 to 0.2816 among different autotetraploid and diploid materials; the genetic distance between different generations of same species was less, but also shows a certain degree of genetic differences. The inter-generational genetic differences of autotetraploid materials were greater than of the diploid materials, which are 0.1359 to 0.0485 as the genetic distance; comparing with the 0.0388 of diploid materials. The result shows that high generation inbred autotetraploid rice material has a certain genetic stability, but the genetic variation between generations is still strong comparing with the source diploid materials. It indicates that, after many generations of purification cultivation, autotetraploid rice materials established a relatively stable genetic structure, at the same time, stronger variability than its diploid parents are useful in the breeding of new varieties, provides a genetic foundation to the agronomic and quality traits improvement. In addition, the analysis result shows that the through the molecular marker-assisted testing, rice genetic polymorphism between materials can effectively distinguish different species, provides a certain basis for molecular identification of varieties of rice. A series of investigation such as cytological identification, statistical analysis of agronomic traits, molecular marker-assisted cluster analysis was applied in this experiment to research genetic pattern of autotetraploid rice high generation inbred lines, revealed the genetic characteristics and the genetic composition of agronomic traits, provides a theoretical basis for the further selection of high quality autotetraploid rice.
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IEECAS SKLLQG
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IEECAS SKLLQG
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IEECAS SKLLQG