植物营养发育.PPT

第四章 植物胚胎发育 胚发生的起始控制 极性的建立 组织分化概述 胚胎区域类型的形成 胚乳和胚柄的形成和作用 种子大小的决定 The trimethylation of lysine 27 of histone 3 (H3K27me3) is the landmark of Polycomb Repressive Complex2 (PRC2) function and is associated with gene repression. In homozygous null mutants of Arabidopsis PRC2, H3K27me3 is globally lost in these mutants. Surprisingly, initial body plant organization and embryo development is largely independent of PRC2 action, which is in sharp contrast to embryonic lethality of PRC2 mutants in animals. However, PRC2 is required to switch from embryonic to seedling phase, and mutant seeds showed enhanced dormancy and germination defects. Indeed, many genes controlling seed maturation and dormancy are marked by H3K27me3 and are upregulated upon loss of PRC2. the here-discovered key role of PRC2 during the developmental phase transition from embryo to seedling growth reveals the adaptation of conserved molecular mechanisms to carry out new functions. where imprinted gene regulation is essential for embryonic development. However, it seems to be more flexible in plants, as imprinting requirements can be bypassed to allow the development of clonal offspring in apomicts. 在哺乳动物配子发生和胚发生过程中，原来的甲基化完全丢失，由从头合成的Dnmt3a 和 Dnmt3b对完全无甲基化的DNA进行甲基化。DMR1和DMR2与其有同源序列。 met1突变胚中一些参与组织分化形态建成的基因也发生变化，YODA (YDA)表达增加，WUSCHEL-RELATED HOMEOBOX2 (WOX2)和WOX8表达下降。YDA编码MAPKKK，参与胚和胚柄专一特性功能。WOX2和 WOX8分别在发育胚的顶端和基部细胞系专一表达, 编码影响细胞分裂的homeodomain转录因子。 通过影响生长素分布和与组织分化有关基因的表达，基因组DNA甲基化控制胚极性的建立。 生长素的极性分布决定顶端基部极性 生长素运输方向在早期胚发生阶段是从基部向上运输，即从胚柄向原胚方向运输，到32细胞球型胚阶段，生长素运输方向开始改变为向基部运输 。 生长素在胚发生过程中运出细胞的载体在拟南芥是PIN1、PIN3、PIN4 、 PIN7 。PIN7在两细胞阶段基部细胞的顶端膜表达，一直到32细胞阶段在胚柄细胞顶端膜表达，这种表达方式说明生长素运输方向是从基部向上运输。DR5报告基因是在?-葡(萄)糖苷酸酶（?-glucuronidase ，GUS）或绿色荧光蛋白（green fluorescent protein ，GFP）前启动子中加上多个重复的生长素反应因子（Auxin Response Factor,ARF）作用的生长素反应元件（auxin response elements ，auxREs）TGTCTC，报告基因可以被生长素激活，用来反映生长素的存在和作用。在野生型胚中，DR5在顶端细胞表达，但应用生长素抑制剂或在pin7突变体中DR5在基部细胞表达，进一步证明早期胚中生长素从基部向顶部运输 。 Immunolocalization of AtPIN1. The presumptive auxin ef