SPL转录因子调控植物花发育及其分子机制研究进展

SPL(squamosa promoter-binding protein-like)转录因子是植物所特有的一类基因家族,广泛存在于绿色植物中,在植物生长发育中具有重要作用。花发育是植物生殖发育中最为重要的一个过程,涉及不同发育方式的转变,即开花决定、花的发端和花器官发生与发育。简要综述了SPL基因的结构与功能并着重阐述了SPL基因在植物花发育过程中的分子机制及生物学功能。最后总结出: SPL转录因子可直接或间接通过参与光周期途径,赤霉素途径及年龄途径来调控植物的开花时间; SPL基因可通过直接激活下游花分生组织特异基因,如LEAFY(LFY),从而调控植物的成花转变; SPL基因可通过与下游花器官特征基因相互作用来调控花器官及其育性的发育,如调控花序、花柄的长度与外形及花器官的大小; SPL基因可调控植物大小孢子发生及雌雄配子体发育。据拟南芥的相关研究结果,初步构绘出拟南芥开花调控中的分子机制。     

2013-14 科技要闻

中国科学院华南植物园李洁尉等揭示了种子成熟过程的表观遗传调控机制。该研究对通过表观遗传调控改善作物性状、提高作物产量、解决国粮食安全问题具有重要理论意义。研究表明,种子贮藏蛋白基因的表达受许多不同调控因子影响。然而,学界关于它们的作用机理并不清楚。此次,研究人员发现拟南芥组蛋白去乙酰化酶HDA19突变体种子成熟基因可在幼苗中异位表达,这揭示种子发育表观遗传调控机理     

茉莉酸处理对木本油料植物小桐子早期花序芽基因表达的影响

小桐子(Jatropha curcas)是大戟科多年生木本油料植物,其种子含油率较高(30％～40％),是一种潜在的可再生能源植物.然而小桐子雌花较少,种子产量较低,严重限制其推广应用.茉莉酸是一种重要的植物激素,在花发育过程中起着重要作用.为了解茉莉酸在小桐子花发育过程中的作用,应用第二代高通量测序方法,对1.0 mmol/L茉莉酸处理24 h后的小桐子早期花序芽进行转录组测序分析,观测茉莉酸对花发育及相关基因转录的影响.结果显示,外源茉莉酸处理导致早期花序芽中1259个基因上调表达,695个基因下调表达;其中,10个与成花转变相关,8个与花器官发育相关以及18个与茉莉酸合成和信号转导途径相关的拟南芥同源基因表达发生了显著变化,但是花器官表型并未有显著的改变.差异表达基因的GO注释显示,"响应茉莉酸"功能分类基因在上调表达基因群中富集,表明在小桐子中存在可以响应外源茉莉酸处理的信号途径.茉莉酸处理后,与拟南芥花器官发育相关基因同源JcFUL、JcSRS、JcSEP1、JcAGL61、JcWOX1、JcTPR4和JcSEU下调表达,然而小桐子花器官表型没有明显改变,说明在小桐子中这些基因的变化表达不足以改变小桐子花器官表型.实验结果对解析茉莉酸在小桐子花发育调控过程中的作用有一定的参考价值.     

拟南芥基因转录调控关系的计算方法研究

拟南芥是一种重要的模式植物,已被广泛应用于植物生物学研究.基于基因表达谱和序列信息构建了预测拟南芥基因调控关系的数学模型.通过支持向量机和夹克刀的测试,结果表明该方法在拟南芥基因调控关系的预测工作中有很好的表现.利用计算方法预测拟南芥基因转录调控关系可为实验室研究提供一定理论依据.     

甘蓝型油菜角果和种子发育相关的Bna-novel-miR432功能分析

为了探究miRNA在油菜花器官发育、角果和种子发育中的功能，本研究在甘蓝型油菜不同发育阶段角果和种子小RNA测序数据分析中，筛选到一个新的miRNA，命名为Bna-novel-miR432，并将其转化至拟南芥中进行功能分析.研究发现过表达Bna-novel-miR432拟南芥开花时间显著提前，并且发现其雄蕊发育存在缺陷，导致雌蕊授粉不完全. 在对其花粉育性进行检测时发现过表达材料花粉育性显著降低. 对角果发育统计发现，过表达拟南芥株系的角果变短，角果中种子数量显著减少，且部分角果存在败育的现象. 此外，在种子发育方面，过表达材料种子中胚的发育也出现滞后的现象. 以上结果表明，Bna-novel-miR432参与调控植物开花时间、雄蕊和花粉的发育以及果实的形成等生物学过程中发挥一定的调控功能.     

赤霉素关键蛋白DELLA对植物雄蕊发育研究进展

在植物生长发育过程中,DELLA蛋白作为响应赤霉素（GA）信号通路负调控因子,其家族成员在植物雄蕊组织中均有表达,对植物生长发育起着重要作用.通过近几年研究发现,无论是拟南芥还是水稻,DELLA基因的突变都会导致植株的雄性不育.该文综述了DELLA蛋白基本特征、参与赤霉素信号通路的调控,以及在雄蕊发育中的作用,为今后的研究提供一定的理论基础.     

植物抗寒基因工程研究最新进展

低温是限制农作物分布和产量的一种非生物胁迫因素，因此提高植物的抗寒性对农业具有十分重要的意义。许多植物在经过一段时间的非冻低温作用后其抗寒性获得增强，这种现象被称为植物的低温驯化。最近10年来，随着对植物低温驯化的分子机制研究的不断深入，植物抗寒基因工程研究获得了长足的进展。目前应用于植物抗寒基因工程的基因包括两大类：保护基因与调控基因，两者都表现出良好的应用前景。然而，植物抗寒基因工程研究领域也存在着大量的问题急需解决。     

YUCCA基因家族在拟南芥胚胎发育过程中的表达模式研究

生长素作为重要的植物激素,在植物胚胎的形态建成和生长发育过程中起到关键的调控作用.依赖色氨酸的IPA途径是模式植物拟南芥中生长素合成的主要途径.以色氨酸为前体合成的IPA在YUCCA(YUC)的催化下生成IAA,这一过程也是生长素生物合成的限速步骤.拟南芥有11个YUC编码基因,目前对这11个YUC基因在植物胚胎发育过程中的表达模式还没有系统的研究.本文以Pro YUC:GUS(YUC1:GUS、YUC2:GUS、YUC4:GUS、YUC6:GUS、YUC8:GUS和YUC9:GUS)转基因植株为实验材料,观察在胚胎发育不同时期YUC基因的表达情况.结果表明:YUC1、YUC2和YUC4基因在拟南芥胚胎发育的后期(鱼雷期、拐杖期和成熟期)有极其微弱的表达;YUC6基因在拟南芥胚胎发育过程中没有表达;YUC8和YUC9在胚发育过程中有非常强的表达,其中YUC8主要在胚胎发育的三角胚时期、心形期和鱼雷期表达,而且在整个胚中表达,在拐杖期胚和成熟胚中YUC8特异在下胚轴和胚根中表达;YUC9在胚胎发育的球形期、三角胚时期和心形期有较强的表达,在胚发育的后期则在胚根有微弱的表达.这些结果表明拟南芥胚胎发育过程中存在生长素的本地合成,而且YUC8和YUC9是胚胎中参与生长素合成的主要基因.     

龙眼DlIKU1基因的克隆与功能分析

拟南芥(Arabidopsis thaliana)IKU1基因突变可以产生较小的种子.根据植物IKU1同源基因的高度保守性,设计引物,PCR扩增获得龙眼(Dimocarpus longan Lour.)同源基因Dl IKU1的编码序列,其编码的蛋白含有VQ保守基序,与其他物种中同源蛋白的氨基酸序列存在较高的同源性;由于龙眼遗传转化的限制,构建Dl IKU1基因的超量表达载体并转化拟南芥iku1突变体,统计转基因植株子代种子的长宽变化,结果表明:龙眼Dl IKU1基因的超量表达可以显著增加拟南芥种子的大小,说明龙眼Dl IKU1基因可以影响种子发育.上述结果为研究龙眼种子发育机理提供了理论和实验依据,有助于龙眼产业的发展.     

半月科技要闻

<正>发现水稻籽粒大小关键调控基因近日,中国科学院院士、华中农业大学张启发课题组在谷粒大小和粒型的调控研究方面取得重大进展:研究证实了水稻中GS3基因控制水稻籽粒大小,发现了该基因中     

Bypassing genomic imprinting allows seed development

In developing progeny of mammals the two parental genomes are differentially expressed according to imprinting marks, and embryos with only a uniparental genetic contribution die. Gene expression that is dependent on the parent of origin has also been observed in the offspring of flowering plants, and mutations in the imprinting machinery lead to embryonic lethality, primarily affecting the development of the endosperm-a structure in the seed that nourishes the embryo, analogous to the function of the mammalian placenta. Here we have generated Arabidopsis thaliana seeds in which the endosperm is of uniparental, that is, maternal, origin. We demonstrate that imprinting in developing seeds can be bypassed and viable albeit smaller seedlings can develop from seeds lacking a paternal contribution to the endosperm. Bypassing is only possible if the mother is mutant for any of the FIS-class genes, which encode Polycomb group chromatin-modifying factors. Thus, these data provide functional evidence that the action of the FIS complex balances the contribution of the paternal genome. As flowering plants have evolved a special reproduction system with a parallel fusion of two female with two male gametes, our findings support the hypothesis that only with the evolution of double fertilization did the action of the FIS genes become a requirement for seed development. Furthermore, our data argue for a gametophytic origin of endosperm in flowering plants, thereby supporting a hypothesis raised in 1900 by Eduard Strasburger.     

SHATTERPROOF MADS-box genes control seed dispersal in Arabidopsis

The fruit, which mediates the maturation and dispersal of seeds, is a complex structure unique to flowering plants. Seed dispersal in plants such as Arabidopsis occurs by a process called fruit dehiscence, or pod shatter. Few studies have focused on identifying genes that regulate this process, in spite of the agronomic value of controlling seed dispersal in crop plants such as canola. Here we show that the closely related SHATTERPROOF (SHP1) and SHATTERPROOF2 (SHP2) MADS-box genes are required for fruit dehiscence in Arabidopsis. Moreover, SHP1 and SHP2 are functionally redundant, as neither single mutant displays a novel phenotype. Our studies of shp1 shp2 fruit, and of plants constitutively expressing SHP1 and SHP2, show that these two genes control dehiscence zone differentiation and promote the lignification of adjacent cells. Our results indicate that further analysis of the molecular events underlying fruit dehiscence may allow genetic manipulation of pod shatter in crop plants.     

Plant retinoblastoma homologues control nuclear proliferation in the female gametophyte

Haploid spores of plants divide mitotically to form multicellular gametophytes. The female spore (megaspore) of most flowering plants develops by means of a well-defined programme into the mature megagametophyte consisting of the egg apparatus and a central cell. We investigated the role of the Arabidopsis retinoblastoma protein homologue and its function as a negative regulator of cell proliferation during megagametophyte development. Here we show that three mutant alleles of the gene for the Arabidopsis retinoblastoma-related protein, RBR1 (ref. 4), are gametophytic lethal. In heterozygous plants 50% of the ovules are aborted when the mutant allele is maternally inherited. The mature unfertilized mutant megagametophyte fails to arrest mitosis and undergoes excessive nuclear proliferation in the embryo sac. Supernumerary nuclei are present at the micropylar end of the megagametophyte, which develops into the egg apparatus and central cell. The central cell nucleus, which gives rise to the endosperm after fertilization, initiates autonomous endosperm development reminiscent of fertilization-independent seed (fis) mutants. Thus, RBR1 has a novel and previously unrecognized function in cell cycle control during gametogenesis and in the repression of autonomous endosperm development.     

Delayed activation of the paternal genome during seed development

Little is known about the timing of the maternal-to-zygotic transition during seed development in flowering plants. Because plant embryos can develop from somatic cells or microspores, maternal contributions are not considered to be crucial in early embryogensis. Early-acting embryo-lethal mutants in Arabidopsis, including emb30/gnom which affects the first zygotic division, have fuelled the perception that both maternal and paternal genomes are active immediately after fertilization. Here we show that none of the paternally inherited alleles of 20 loci that we tested is expressed during early seed development in Arabidopsis. For genes that are expressed at later stages, the paternally inherited allele becomes active three to four days after fertilization. The genes that we tested are involved in various processes and distributed throughout the genome, indicating that most, if not all, of the paternal genome may be initially silenced. Our findings are corroborated by genetic studies showing that emb30/gnom has a maternal-effect phenotype that is paternally rescuable in addition to its zygotic lethality. Thus, contrary to previous interpretations, early embryo and endosperm development are mainly under maternal control.     

Identification of diploid endosperm in an early angiosperm lineage

In flowering plants, the developmental and genetic basis for the establishment of an embryo-nourishing tissue differs from all other lineages of seed plants. Among extant nonflowering seed plants (conifers, cycads, Ginkgo, Gnetales), a maternally derived haploid tissue (female gametophyte) is responsible for the acquisition of nutrients from the maternal diploid plant, and the ultimate provisioning of the embryo. In flowering plants, a second fertilization event, contemporaneous with the fusion of sperm and egg to yield a zygote, initiates a genetically biparental and typically triploid embryo-nourishing tissue called endosperm. For over a century, triploid biparental endosperm has been viewed as the ancestral condition in extant flowering plants. Here we report diploid biparental endosperm in Nuphar polysepalum, a basal angiosperm. We show that diploid endosperms are common among early angiosperm lineages and may represent the ancestral condition among flowering plants. If diploid endosperm is plesiomorphic, the triploid endosperms of the vast majority of flowering plants must have evolved from a diploid condition through the developmental modification of the unique fertilization process that initiates endosperm.     

The development of embryo and endosperm inRanalisma rostratum

The embryo and endosperm development inRanalisma rostratum was studied in this paper. The zygote divides by a transverse wall to form basal and apical cells. The larger basal cell undergoes no further division and becomes the basal suspensor cell. The embryo development conforms to the Caryophyllad type. In the mature seed, the embryo is U-shaped and forms the embryonic shoot apex accompanied by one intraseminal leaf. The endosperm development belongs to the Hebolial type. The primary endosperm nucleus (invariably lying in the chalazal part of the embryo sac) divides and forms a large micropylar chamber and a small chalazal chamber. The chalazal endosperm chamber remains uninucleate, and free nuclear divisions occur in the micropylar chamber. Cellularization of the endosperm begins simultaneously from the micropylar and the chalazal ends. The cellularization is related to phragmoplast and cell plate, and does not arise from the free growth wall. Accompanying the embryo development, the endosperm degrades and does not exist in the mature seed. In the proembryo developing stages, the basal suspensor cell and chalazal endosperm cell grow to a considerable size. The nucleus and cytoplasm stain densely. Sometimes, binucleate or trinucleate was observed in both of the two cells. In the laterally concave embryo stage, they begin to degenerate which coincides with the cellularization of the endosperm. They are presumed to participate in the nourishment of the embryo proper in the early stages of embryo development. Supported by the National Natural Science Foundation of China Wang Jianbo: born in May. 1964, Ph. D.     

Gamete formation without meiosis in Arabidopsis

Apomixis, the formation of asexual seeds in plants, leads to populations that are genetically uniform maternal clones. The transfer of apomixis to crop plants holds great promise in plant breeding for fixation of heterozygosity and hybrid vigour because it would allow the propagation of hybrids over successive generations. Apomixis involves the production of unreduced (diploid) female gametes that retain the genotype of the parent plant (apomeiosis), followed by parthenogenetic development of the egg cell into an embryo and the formation of functional endosperm. The molecular mechanisms underlying apomixis are unknown. Here we show that mutation of the Arabidopsis gene DYAD/SWITCH1 (SWI1), a regulator of meiotic chromosome organization, leads to apomeiosis. We found that most fertile ovules in dyad plants form seeds that are triploid and that arise from the fertilization of an unreduced female gamete by a haploid male gamete. The unreduced female gametes fully retain parental heterozygosity across the genome, which is characteristic of apomeiosis. Our results show that the alteration of a single gene in a sexual plant can bring about functional apomeiosis, a major component of apomixis.     

水稻和小麦胚乳发育的比较

以水稻盐粳235和小麦杨麦5号为,和整体解剖和树脂包埋切片等方法比较研究了这两种作物胚乳细胞分裂、分化和充实的特点,两者发育的主要差异有：（1）水稻胚囊呈香蕉形;小麦胚乳囊呈现瓜子形,小麦的游离核与胚乳细胞要比水稻大;（2）水稻游离核的分裂以无丝分裂为主,而小麦游离核的分裂以有丝分裂为主,水稻游离核及细胞的分裂速度较小麦快,（3）水稻胚乳细胞中淀粉体约在花后第4天出现,蛋白质体约在花后第5天出现;小麦胚乳细胞中淀粉体约在花后第7天出现,而蛋白质体约在花后第9天出现;（4）水稻胚乳淀粉体中含有多个淀粉粒,而小麦胚乳淀粉体中仅含有一个淀粉粒;（5）水稻胚乳含两种蛋白质体类型,即PB1和PB2,而小麦胚乳只含有一种蛋白质体;（6）水稻胚乳的背部有多层糊粉层细胞,其细胞壁上没有内突,而小麦腹部（沟）维管束的糊粉层细胞壁上有内,帝些细胞进而转化为胚乳转移细胞。     

四川无融合水稻（SAR—1）研究

四川无融合水稻(SAR-1)是1988年春在南繁育种材料中首次发现的新种质.该材料花粉高度不育,能大量单性结实.其基因不仅能世代相传,而且可通过杂交转移到其他材料上.经细胞胚胎学研究表明:SAR-1具孤雌生殖特性,还可能具有不定胚发生途径.SAR-1的极核未经受精,能自发形成胚乳.