胚乳细胞壁的建成及其与胞质微管的关系

综述了被子植物核型胚乳发育的不同阶段(合胞体时期、初始垂周壁形成期、初始平周壁形成期和胚乳细胞分裂增生期)胚乳细胞壁建成的各种观点,以及在各个阶段中微管与胚乳细胞壁建成之间关系的最新研究进展。并分别比较了胚乳细胞化过程不同阶段胚乳游离核或胚乳细胞有丝分裂与分生组织细胞有丝分裂微管周期的差异。     

The titan mutants of Arabidopsis are disrupted in mitosis and cell cycle control during seed development

We describe in this report a novel class of mutants that should facilitate the identification of genes required for progression through the mitotic cell cycle during seed development in angiosperms. Three non-allelic titan ( ttn ) mutants with related but distinct phenotypes are characterized. The common feature among these mutants is that endosperm nuclei become greatly enlarged and highly polyploid. The mutant embryo is composed of a few giant cells in ttn1 , several small cells in ttn2 , and produces a normal plant in ttn3 . Condensed chromosomes arrested at prophase of mitosis are found in the free nuclear endosperm of ttn1 and ttn2 seeds. Large mitotic figures with excessive numbers of chromosomes are visible in ttn3 endosperm. The ttn1 mutation appears to disrupt cytoskeletal organization because endosperm nuclei fail to migrate to the chalazal end of the seed. How double fertilization leads to the establishment of distinct patterns of mitosis and cytokinesis in the embryo and endosperm is a central question in plant reproductive biology. Molecular isolation of TITAN genes should help to answer this question, as well as related issues concerning cell cycle regulation, chromosome movement and endosperm identity in angiosperms.     

D-type cyclins control cell division and developmental rate during Arabidopsis seed development

Seed development in Arabidopsis is characterized by stereotypical division patterns, suggesting that coordinated control of cell cycle may be required for correct patterning and growth of the embryo and endosperm. D-type cyclins (CYCD) are key cell cycle regulators with roles in developmental processes, but knowledge regarding their involvement in seed development remains limited. Here, a family-wide gene expression, and loss- and gain-of-function approach was adopted to reveal additional functions for CYCDs in the development of seed tissues. CYCD genes have both discrete and overlapping tissue-specific expression patterns in the seed as revealed by GUS reporter gene expression. Analysis of different mutant combinations revealed that correct CYCD levels are required in seed development. The CYCD3 subgroup is specifically required as its loss caused delayed development, whereas overexpression in the embryo and endosperm of CYCD3;1 or a previously uncharacterized gene, CYCD7;1, variously leads to induced proliferation, abnormal phenotypes, and elevated seed abortion. CYCD3;1 overexpression provoked a delay in embryonic developmental progression and abnormalities including additional divisions of the hypophysis and suspensor, regions where CYCD3 genes are normally expressed, but did not affect endosperm development. Overexpression of CYCD7;1, not normally expressed in seed development, promoted overgrowth of both embryo and endosperm through increased division and cell enlargement. In contrast to post-germination growth, where pattern and organ size is not generally related to division, results suggest that a close control of cell division through regulation of CYCD activity is important during seed development in conferring both developmental rate and correct patterning.     

The female gametophyte and the endosperm control cell proliferation and differentiation of the seed coat in Arabidopsis

Double fertilization of the female gametophyte produces the endosperm and the embryo enclosed in the maternal seed coat. Proper seed communication necessitates exchanges of signals between the zygotic and maternal components of the seed. However, the nature of these interactions remains largely unknown. We show that double fertilization of the Arabidopsis thaliana female gametophyte rapidly triggers sustained cell proliferation in the seed coat. Cell proliferation and differentiation of the seed coat occur in autonomous seeds produced in the absence of fertilization of the multicopy suppressor of ira1 (msi1) mutant. As msi1 autonomous seeds mostly contain autonomous endosperm, our results indicate that the developing endosperm is sufficient to enhance cell proliferation and differentiation in the seed coat. We analyze the effect of autonomous proliferation in the retinoblastoma-related1 (rbr1) female gametophyte on seed coat development. In contrast with msi1, supernumerary nuclei in rbr1 female gametophytes originate mainly from the endosperm precursor lineage but do not express an endosperm fate marker. In addition, defects of the rbr1 female gametophyte also reduce cell proliferation in the ovule integuments before fertilization and prevent further differentiation of the seed coat. Our data suggest that coordinated development of the seed components relies on interactions before fertilization between the female gametophyte and the surrounding maternal ovule integuments and after fertilization between the endosperm and the seed coat.     

Embryological Studies of Paphiopedilum godefroyae Stein

P. godefroyae is one of the diandrous species of rather primitive orchids. The cytokinesis of PMCs conforms to simultaneous type. The arrangement of microspores in a tetrad is tetrahedral or isobilateral. The first mitosis in a pollen grain is unequal and results in the formation of two unequal cells. The small one is the generative cell and the large one, the vegetative cell. The wall material between them is callose which is easily detectable under the fluorescence microscope. When the generative cell detaches from the microspore wall and migrates into the cytoplasm of the vegetative cell, the callose wall disappears and a thin PAS-positive wall Was observed around the generative cell. The PAS-positive wall remains untill anthesis. The tapetum is of secretory type and its cells are binucleate. With the degradation of the tapetal cells, they discharge a lot of yellow, amorphous, sticky mass into the pollen sac. The pollens distribute in it to form a sticky pollen mass. The ovule has single integument and one layered nucellus around the magaspore mother cell. The mature embryo sac consists of eight or six cells and conforms to the Allium type. The interval between pollination and fertilization is about 45 days and the normal double fertilization has been observed. The primary endosperm cell undergoes one division only and results in the formation of 2 nucleate endosperm. The dormancy period of zygote lasts 45–50 days. During the development of the embryo, a suspensor consisting of a row of two to four cells is formed. It takes more than six months from the pollination to the maturation of the seed. The embryo in the mature seed is just an ellipsoidal mass of 120–140 cells without differentiation. The endosperm and suspensor are all degenerated in the mature seed.     

Genetic analysis as a tool to investigate the molecular mechanisms underlying seed development in maize

BACKGROUND: In angiosperms the seed is the outcome of double fertilization, a process leading to the formation of the embryo and the endosperm. The development of the two seed compartments goes through three main phases: polarization, differentiation of the main tissues and organs and maturation. SCOPE: This review focuses on the maize kernel as a model system for developmental and genetic studies of seed development in angiosperms. An overview of what is known about the genetic and molecular aspects underlying embryo and endosperm formation and maturation is presented. The role played by embryonic meristems in laying down the plant architecture is discussed. The acquisition of the different endosperm domains are presented together with the use of molecular markers available for the detection of these domains. Finally the role of programmed cell death in embryo and endosperm development is considered. CONCLUSIONS: The sequence of events occurring in the developing maize seed appears to be strictly regulated. Proper seed development requires the co-ordinated expression of embryo and endosperm genes and relies on the interaction between the two seed components and between the seed and the maternal tissues. Mutant analysis is instrumental in unravelling the genetic control underlying the formation of each compartment as well as the molecular signals interplaying between the two compartments.     

ANAPHASE PROMOTING COMPLEX/CYCLOSOME‐mediated cyclin B1 degradation is critical for cell cycle synchronization in syncytial endosperms

Although it is known that in most angiosperms mitosis in early endosperm development is syncytial and synchronized,it is unclear how the synchronization is regulated.We showed previously that APC11,also named ZYG1,in Arabidopsis activates zygote division by interaction and degradation of cyclin B1.Here,we report that the mutation in APC11/ZYG1 led to unsynchronized mitosis and over-accumulation of cyclin B1-GUS in the endosperm.Mutations in two other APC subunits showed similar defects.Transgenic expression of stable cyclin B1 in the endosperm also caused unsynchronized mitosis.Further,downregulation of APC11 generated multi-nucleate somatic cells with unsynchronized mitotic division.Together,our results suggest that APC/C-mediated cyclin B1 degradation is critical for cell cycle synchronization.     

Sexual and apomictic reproduction in Hieracium subgenus pilosella are closely interrelated developmental pathways

Seed formation in flowering plants requires meiosis of the megaspore mother cell (MMC) inside the ovule, selection of a megaspore that undergoes mitosis to form an embryo sac, and double fertilization to initiate embryo and endosperm formation. During apomixis, or asexual seed formation, in Hieracium ovules, a somatic aposporous initial (AI) cell divides to form a structurally variable aposporous embryo sac and embryo. This entire process, including endosperm development, is fertilization independent. Introduction of reproductive tissue marker genes into sexual and apomictic Hieracium showed that AI cells do not express a MMC marker. Spatial and temporal gene expression patterns of other introduced genes were conserved commencing with the first nuclear division of the AI cell in apomicts and the mitotic initiation of embryo sac formation in sexual plants. Conservation in expression patterns also occurred during embryo and endosperm development, indicating that sexuality and apomixis are interrelated pathways that share regulatory components. The induction of a modified sexual reproduction program in AI cells may enable the manifestation of apomixis in HIERACIUM:     

The globby1-1 (glo1-1) mutation disrupts nuclear and cell division in the developing maize seed causing alterations in endosperm cell fate and tissue differentiation

Cereal endosperm tissues account for most of the world's calorific intake, yet the regulation of monocot seed development remains poorly understood. The maize endosperm originates with a series of free-nuclear divisions, followed by cellularisation and subsequent formation of a range of functional cellular domains. We describe the isolation and characterisation of a mutation that induces aberrant globular embryo and endosperm morphology, globby1-1 (glo1-1). Our data indicate that glo1-1 plays a role in nuclear division and cytokinesis in the developing seed. Pattern formation in the embryo is severely impaired with development arresting at premature stages, while in the endosperm, the effects of the glo1-1 mutation are manifest at the free-nuclear or syncytial stage. During cellularisation, and at later stages of development, aberrant cell division and localised domains of cell proliferation are apparent in glo1-1 endosperms. As a consequence, cell fate acquisition and subsequent differentiation of endosperm tissues are affected to varying degrees of severity. To date, it has been hypothesised that BETL cell fate is specified in the syncytium and that cell files subsequently develop in response to a gradient of signal(s) derived from the maternal pedicel region. Based on our findings, however, we propose that specification of BETL cells is an irreversible event that occurs within a narrow window of syncytial development, and that BETL cell identity is subsequently inherited in a lineage-dependent manner. Additionally, our data suggest that acquisition of aleurone cell fate does not solely rely upon signalling from the maternal surrounding tissue to the periphery of the endosperm, as previously thought, but that other factor(s) present within the endosperm are involved.     

Effects of APETALA2 on embryo, endosperm, and seed coat development determine seed size in Arabidopsis

Arabidopsis APETALA2 (AP2) controls seed mass maternally, with ap2 mutants producing larger seeds than wild type. Here, we show that AP2 influences development of the three major seed compartments: embryo, endosperm, and seed coat. AP2 appears to have a significant effect on endosperm development. ap2 mutant seeds undergo an extended period of rapid endosperm growth early in development relative to wild type. This early expanded growth period in ap2 seeds is associated with delayed endosperm cellularization and overgrowth of the endosperm central vacuole. The subsequent period of moderate endosperm growth is also extended in ap2 seeds largely due to persistent cell divisions at the endosperm periphery. The effect of AP2 on endosperm development is mediated by different mechanisms than parent-of-origin effects on seed size observed in interploidy crosses. Seed coat development is affected; integument cells of ap2 mutants are more elongated than wild type. We conclude that endosperm overgrowth and/or integument cell elongation create a larger postfertilization embryo sac into which the ap2 embryo can grow. Morphological development of the embryo is initially delayed in ap2 compared with wild-type seeds, but ap2 embryos become larger than wild type after the bent-cotyledon stage of development. ap2 embryos are able to fill the enlarged postfertilization embryo sac, because they undergo extended periods of cell proliferation and seed filling. We discuss potential mechanisms by which maternally acting AP2 influences development of the zygotic embryo and endosperm to repress seed size.     

Maternal control of integument cell elongation and zygotic control of endosperm growth are coordinated to determine seed size in Arabidopsis

We use Arabidopsis thaliana as a model to investigate coordination of cell proliferation and cell elongation in the three components that develop side by side in the seed. Two of these, the embryo and its nurturing annex, the endosperm, are placed under zygotic control and develop within the seed integument placed under maternal control. We show that integument cell proliferation and endosperm growth are largely independent from each other. By contrast, prevention of cell elongation in the integument by the mutation transparent testa glabra2 (ttg2) restricts endosperm and seed growth. Conversely, endosperm growth controlled by the HAIKU (IKU) genetic pathway modulates integument cell elongation. Combinations of TTG2 defective seed integument with reduction of endosperm size by iku mutations identify integument cell elongation and endosperm growth as the primary regulators of seed size. Our results strongly suggest that a cross talk between maternal and zygotic controls represents the primary regulator of the coordinated control of seed size in Arabidopsis.     

Endosperm development

There is a renewal of interest in endosperm development. Recent studies are leading the way to a better understanding of fundamental processes such as cell cycle control and the mechanisms of imprinting. A more global view of interactions between the endosperm and the embryo is emerging and will initiate an integrated approach to the study of seed development.     

A Comparison of Leek (Allium porrum) and Onion (Allium cepa) Seed Development

Leek and onion seed dry weight increased exponentially for thefirst 31 days after flowering (DAF) but thereafter the increasein dry weight was slower. Before maximum seed dry weight wasreached at 45 DAF in onion and 59 to 66 DAF in leek, seed moisturecontent, seed oxygen uptake and conductivity of the seed steepwater fell from initially high levels. Although some seeds germinated31 DAF in both species, full germination in both was not reacheduntil 66–80 DAF. Tolerance of the seed to artificial dryingimmediately after harvesting occurred 45 DAF in onion and 74DAF in leek. Free nuclear division continued in the endospermuntil 17–22 DAF in onion and until 31–35 DAF inleek but it was not until 45 DAF in onion and 66 DAF in leekthat the embryo and endosperm filled the cavity formed by thepericarp. After formation of cell walls in the endosperm thepattern of change in cell number in both species was similar.The shrunken appearance of the seed coat in leek, which occurredearly in seed development, was associated with the period offree nuclear division in the endosperm and, in addition, thepericarp was thinner than in onion. There was no evidence thatthe shrunken seed coat early in development was associated withself as opposed to open-pollination. Allium porrum, Allium cepa, seed development, endosperm, embryo, cell number, germination, respiration, seed leachates     

Dynamic analyses of the expression of the HISTONE::YFP fusion protein in arabidopsis show that syncytial endosperm is divided in mitotic domains

During early seed development, nuclear divisions in the endosperm are not followed by cell division, leading to the development of a syncytium. The simple organization of the Arabidopsis endosperm provides a model in which to study the regulation of the cell cycle in relation to development. To monitor nuclear divisions, we constructed a HISTONE 2B::YELLOW FLUORESCENT PROTEIN gene fusion (H2B::YFP). To validate its use as a vital marker for chromatin in plants, H2B::YFP was expressed constitutively in Arabidopsis. This enabled the observation of mitoses in living root meristems. H2B::YFP was expressed specifically in Arabidopsis syncytial endosperm by using GAL4 transactivation. Monitoring mitotic activity in living syncytial endosperm showed that the syncytium was organized into three domains in which nuclei divide simultaneously with a specific time course. Each mitotic domain has a distinct spatiotemporal pattern of mitotic CYCLIN B1;1 accumulation. The polar spatial organization of the three mitotic domains suggests interactions between developmental mechanisms and the regulation of the cell cycle.