水蔗草兼性无融合生殖胚胎学研究

对水蔗草 (ApludamuticaL .)的生殖方式进行研究 ,结果表明水蔗草进行兼性无融合生殖。胚囊发育分为两种类型 ,即有性生殖的蓼型和无孢子生殖的大黍型。无融合生殖胚囊频率为 6 0 .74%。在大孢子母细胞发育至四分体后 ,珠孔端的 3个大孢子解体。合点端的大孢子未解体时 ,邻近大孢子的 1个珠心细胞开始特化 ,形成无融合生殖的原始细胞 ,由该原始细胞发育形成有 1个卵细胞、1个助细胞和 2个极核的四核胚囊。     

水蔗草兼性无融合生殖胚胎学研究

对水蔗草(Apluda mutica L.)的生殖方式进行研究,结果表明水蔗草进行兼性无融合生殖.胚囊发育分为两种类型,即有性生殖的蓼型和无孢子生殖的大黍型.无融合生殖胚囊频率为60.74%.在大孢子母细胞发育至四分体后,珠孔端的3个大孢子解体.合点端的大孢子未解体时,邻近大孢子的1个珠心细胞开始特化,形成无融合生殖的原始细胞,由该原始细胞发育形成有1个卵细胞、1个助细胞和2个极核的四核胚囊.     

四倍体双穗雀稗兼性无孢子生殖的研究

研究了四倍体双穗雀稗(Paspalum distichum L)无孢子生殖胚囊、胚胎发育以及假受精特点。当其大孢子母细胞发育至四分体阶段时,大多数情况下会发生四分体退化,同时有多个特化珠心细胞发育为1—3个无孢子生殖胚囊的现象。成熟无孢子生殖胚囊一般3核,包括1个卵细胞和2个极核。卵细胞在抽穗前就能自发分裂形成原胚团,而极核则在抽穗和传粉后参与假受精形成胚乳。当胚珠内存在多个无孢子生殖胚囊时,只是靠近珠孔端的1个无孢子生殖胚囊内的极核与精核结合,而其它的并不参与。种子成熟后出现很低频率的二胚苗。此外,还能观察到少量的有性生殖胚囊的发育以及有性生殖胚囊和无孢子生殖胚囊在同一胚珠中的发育现象,因此判断该类群为兼性无孢子生殖体。     

雾灵山草地早熟禾多胚囊和多胚现象的研究

无融合生殖是指未经精卵融合而产生后代的特殊生殖方式,它可以分为单倍体无融合生殖和二倍体无融合生殖;对于作物改良意义更大的是二倍体无融合生殖。多胚囊和多胚现象SHI是无融合生殖的表现形式。本文运用石蜡切片法、子房整体透明法研究了雾灵山草地早熟禾〖WTBX〗（Poa pratensis〖WTBZ〗 L.）多胚囊和多胚现象。结果表明,（1）草地早熟禾多胚囊来源有两种：一是来自大孢子母细胞,二是来自珠心细胞;（2）草地早熟禾多胚来源有四个：其一是有性生殖胚,其二是孤雌生殖胚,其三是无配子生殖胚,其四是珠心胚。     

水蔗草胚珠附器的研究

本文对水蔗草的胚珠附器进行研究,结果表明：在功能大孢子时期,珠孔端的1—3个珠心细胞开始特化,发育成胚珠附器;胚珠附器发生时,有些胚珠同时出现无孢子生殖原始细胞;有性生殖和无孢子生殖的胚囊中均有胚珠附器存在;但在无孢子生殖的胚囊中,胚珠附器一般很大,长约是宽的1—3倍;而有性生殖胚囊的胚珠附器的长约是宽的1—2倍;和有性生殖胚囊相比,无孢子生殖胚囊的胚珠附器更加发达;存在发达的胚珠附器是水蔗草无孢子生殖胚囊的特点之一。     

水蔗草胚珠附器的研究

本文对水蔗草的胚珠附器进行研究,结果表明：在功能大孢子时期,珠孔端的1～3个珠心细胞开始特化,发育成胚珠附器;胚珠附器发生时,有些胚珠同时出现无孢子生殖原始细 胞;有性生殖和无孢子生殖的胚囊中均有胚珠附器存在;但在无孢子生殖的胚囊中,胚珠附器一般很大,长约是宽的1～３倍;而有性生殖胚囊的胚珠附器的长约是宽的1～２倍;和有性生殖胚囊相比,无孢子生殖胚囊的胚珠附器更加发达;存在发达的胚珠附器是水蔗草无孢子生殖胚囊的特点之一。     

高粱SSA-1无融合生殖胚胎学研究

经常规石蜡切片法,在光学水平观察了高粱（Ｓｏｒｇｈｕｍ ｂｉｃｏｌｏｒ （Ｌ．） Ｍｏｅｎｃｈ） ＳＳＡ－１ 无融合生殖的胚胎发生。高粱ＳＳＡ－１ 的无融合生殖为无孢子生殖和二倍体孢子生殖两种类型。两种生殖类型的单核胚囊经３ 次有丝分裂形成７ 细胞（８ 核）的成熟胚囊,由卵细胞、２ 个助细胞、２ 个极核和３ 个反足细胞组成。反足细胞迅速分裂增殖,形成由２０—３０ 个细胞组成的细胞团。此外,还具有一定频率的无孢子生殖多孢原和多胚囊现象。在未授粉情况下,卵细胞自发分裂形成典型的禾本科类型单子叶胚。经切片统计表明,ＳＳＡ－１ 的无融合生殖频率为４２％ ,证明该系为一兼性无融合生殖系。文中还讨论了ＳＳＡ－１ 无融合生殖过程的特点。     

外来入侵植物胜红蓟的胚胎学观察及繁殖系统研究

以采自我国广东江门和广州两个种群的胜红蓟(Ageratum conyzoides L.)种子为材料,采用流式细胞种子筛选技术(FCSS)和人工控制授粉实验,对胜红蓟的繁殖系统进行研究,并结合整体透明技术和微分干涉差(DIC)显微镜观察法,对其胚珠发育过程和花药结构进行细胞胚胎学观察。种子筛选结果显示,胜红蓟的种子既可以通过有性生殖产生,又可以通过不需要假受精的无融合生殖产生,属于兼性无融合生殖类型。开放性授粉和套袋处理之间的结实率均较高,分别为88%±1.2%和86.2%±1.2%,两者之间无显著差异;而去雄处理的结实率和开放性授粉、套袋这两种处理之间差异显著。胚珠的细胞胚胎学观察结果发现,胜红蓟的有性生殖胚囊发育方式为蓼型,无融合生殖胚囊的发育方式为山柳菊型。胜红蓟的花粉粒在花药内就开始萌发出花粉管,具有闭花受精特性。研究结果表明闭花受精和兼性无融合生殖等繁殖特性保证了胜红蓟在各种生存环境下的结实量,提高其在新生境中归化和入侵的可能性。     

雾灵山草地早熟禾多胚囊和多胚的研究

无融合生殖是指未经炷卵事例而产生后代的特殊生殖方式,它可以分为单倍体无融合生殖和二倍体无融合生殖;对于作物意义更大的是二倍体无融合生殖。多胚囊和多胚现象ＳＨＩ是无融合生殖的表现形式。本文运用石蜡切片法、子房整体透明法研究了雾灵山草地早熟禾（Ｐｏａ ｐｒａｔｅｎｓｉｓ Ｌ．）多胚囊和多胚现象。结果表明,（１）草地早熟禾多胚囊来源有两种：一是自大孢子母细胞,二是来自珠心细胞;（２）草地早熟禾多 来源有     

无融合生殖在植物育种中的应用

无融合生殖(apomixis)是一种代替有性生殖的,不发生核融合而产生种子的生殖方式。它是介于有性生殖与无性生殖之间的一种特殊生殖方式。     

Dynamics of callose deposition and beta-1,3-glucanase expression during reproductive events in sexual and apomictic Hieracium

Callose accumulates in the walls of cells undergoing megasporogenesis during embryo sac formation in angiosperm ovules. Deficiencies in callose deposition have been observed in apomictic plants and causal linkages between altered callose deposition and apomictic initiation proposed. In apomictic Hieracium, embryo sacs initiate by sexual and apomictic processes within an ovule, but sexual development terminates in successful apomicts. Callose deposition and the events that lead to sexual termination were examined in different Hieracium apomicts that form initials pre- and post-meiosis. In apomictic plants, callose was not detected in initial cell walls and deficiencies in callose deposition were not observed in cells undergoing megasporogenesis. Multiple initial formation pre-meiosis resulted in physical distortion of cells undergoing megasporogenesis, persistence of callose and termination of the sexual pathway. In apomictic plants, callose persistence did not correlate with altered spatial or temporal expression of a β-1,3-glucanase gene (HpGluc) encoding a putative callose-degrading enzyme. Expression analysis indicated HpGluc might function during ovule growth and embryo sac expansion in addition to callose dissolution in sexual and apomictic plants. Initial formation pre-meiosis might therefore limit the access of HpGluc protein to callose substrate while the expansion of aposporous embryo sacs is promoted. Callose deposition and dissolution during megasporogenesis were unaffected when initials formed post-meiosis, indicating other events cause sexual termination. Apomixis in Hieracium is not caused by changes in callose distribution but by events that lead to initial cell formation. The timing of initial formation can in turn influence callose dissolution. Received: 18 April 2000 / Accepted: 10 July 2000     

Apomixis and sexuality in Paspalum simplex: characterization of the mode of reproduction in segregating progenies by different methods

Segregating progenies of crosses between sexual and apomictic genotypes of Paspalum simplex were analysed for the formation of meiotic versus aposporous embryo sacs, zygotic versus parthenogenetic embryos, and autonomous versus pseudogamous endosperms by using cytoembryological and flow cytometric analyses. Reduced and unreduced 8-nucleated embryo sacs were the final product of female gametophyte development in sexual and aposporous genotypes, respectively. An incomplete penetrance of parthenogenesis was detected in aposporous genotypes. The relative DNA content of endosperm nuclei revealed the normal 2:1 maternal to paternal ratio in sexuals and a 4:1 ratio in apomicts, indicating insensitivity of the apomictic genotypes to endosperm imprinting. Apospory, parthenogenesis and pseudogamy are located on a relatively large linkage group and are inherited together with previously developed molecular markers as a single genetic unit in segregating progenies.     

Mode of reproduction is detected by Parth1 and Sex1 SCAR markers in a wide range of facultative apomictic Kentucky bluegrass varieties

Gametophytic apomixis in Kentucky bluegrass (Poa pratensis L.) involves the parthenogenetic development of unreduced eggs from aposporic embryo sacs. Marker-assisted selection for the mode of reproduction in P. pratensis would avoid costly and time-consuming phenotypic progeny tests. We developed and tested two SCAR primer pairs that are associated with the mode of reproduction in P. pratensis. The SCAR primers identified the apomictic and sexual genotypes among progenies of sexual x apomictic crosses with very low bias. Furthermore, when tested on a wide range of Italian and exotic P. pratensis germplasm, they were able to unequivocally distinguish sexual from apomictic genotypes. This system should, therefore, allow new selection models to be set up in this species.     

Reconstruction of reproductive diversity in Hypericum perforatum L. opens novel strategies to manage apomixis

The mode of reproduction was characterized for 113 accessions of the tetraploid facultative apomictic species Hypericum perforatum using bulked or single mature seeds in the flow cytometric seed screen (FCSS). This screen discriminates several processes of sexual or asexual reproduction based on DNA contents of embryo and endosperm nuclei. Seed formation in H. perforatum proved to be highly polymorphic. Eleven different routes of reproduction were determined. For the first time, individual seeds were identified that originated from two embryo sacs: the endosperm from an aposporous and the embryo from the legitimate meiotic embryo sac. Moreover, diploid plants were discovered, which apparently reproduce by a hitherto unknown route of seed formation, that is chromosome doubling within aposporous initial cells followed by double fertilization. Although most plants were tetraploid and facultative sexual/apomictic, diploid obligate sexuals and tetraploid obligate apomicts could be selected. Additionally, genotypes were detected which at a high frequency produced embryos either from reduced parthenogenetic or unreduced fertilized egg cells. The endosperm developed most frequently after fertilization of the central cell in aposporous embryo sacs (pseudogamy) but in few cases also autonomously. The genetic control of apomixis appears to be complex in H. perforatum. Basic material was developed for breeding H. perforatum, and strategies are suggested for elucidation of inheritance as well as evolution of apomixis and for molecular approaches of apomixis engineering.     

Sexual reproduction development in apomictic Eulaliopsis binata (Poaceae)

Apomixis is a particular mode of reproduction that allows progeny formation without meiosis and fertilization. Eulaliopsis binata, a tetraploid apomictic species, is widely used for making paper, rope and mats. There is great potential for fixation of heterosis in E. binata due to autonomous endosperm formation in this species. Although most of its embryo sac originates from nucellus cells, termed apospory, we observed sexual reproduction initiation in 86.8 to 96.8% of the ovules, evidenced by callose deposition on the walls of cells undergoing megasporogenesis. However, only 2-3% mature polygonum-type sexual embryo sacs were confirmed by embryological investigation. Callose was not detected on aposporous initial cell walls. The aposporous initial cells differentiated during pre- and post-meiosis of the megaspore mother cell, while the sexual embryo sac degenerated at the megaspore stage. DNA content ratio of embryo and endosperm in some individuals was 2C:3C, based on flow cytometry screening of seed, similar to that of normal sexual seed. These results confirm that apomictic E. binata has conserved sexual reproduction to a certain degree, which may contribute to maintaining genetic diversity. The finding of sexual reproduction in apomictic E. binata could be useful for research on genetic mechanism of apomixis in E. binata.     

Comparative cyto-embryological investigations of sexual and apomictic dandelions (Taraxacum) and their apomictic hybrids

. In the autonomous apomictic Taraxacum officinale (common dandelion), parthenogenetic egg cells develop into embryos and central cells into endosperm without prior fertilisation. Unreduced (2n) megaspores are formed via meiotic diplospory, a nonreductional type of meiosis. In this paper, we describe the normal developmental pathways of sexual and apomictic reproduction and compare these with the development observed in the apomictic hybrids. In sexual diploids, a standard type of megasporogenesis and embryo sac development is synchronised between florets in individual capitula. In contrast, we observed that megasporogenesis and gametogenesis proceeded asynchronously between florets within a single capitulum of natural triploid apomicts. In addition, autonomous endosperm and embryo development initiated independently within individual florets. Parthenogenetic initiation of embryo development in outdoor apomicts was found to be temperature-dependent. Egg cells produced in natural apomicts were not fertilised after pollination with haploid pollen grains although pollen tubes were observed to grow into their embryo sacs. Both reductional and diplosporous megasporogenesis were observed in individual inflorescences of triploid apomictic hybrids. Embryo and endosperm development initiated independently in natural and hybrid apomicts.     

水稻PDER品系的特点和二倍体孢子生殖胚囊的形成和发育

对水稻（ＯｒｙｚａｓａｔｉｖａＬ．）早发生胚ＰＤＥＲ（ｐｒｅ－ｄｅｖｅｌｏｐｅｄｅｍｂｒｙｏｏｆｒｉｃｅ）品系的特点和细胞胚胎学研究表明,ＰＤＥＲ是二倍体植物２ｎ＝２４,约有５０％胚囊的卵细胞未经受精能自行发育形成胚,成熟种子的萌发和生长速度较常规正常水稻快。ＰＤＥＲ的大孢子母细胞经有丝分裂产生未减数的胚囊,即无融合生殖中的二倍体孢子生殖类型。在胚囊形成和发育过程中有如下几个特点：（１）孢原细胞至大孢子母细胞分裂前的过渡期持续时间较长,孢原细胞和大孢子母细胞的细胞质比周围的珠心细胞质稀淡。（２）大孢子母细胞经二次有丝分裂后形成直线排列的三个细胞（三分体）,珠孔端的两个解体,合点端的一个发育为功能细胞,有少数胚囊的三个细胞全部解体形成败育胚囊。（３）功能细胞经三次连续核分裂形成具八核七个细胞的成熟胚囊,它的结构与常规正常水稻基本相同,但助细胞呈长形而没有回抱着卵细胞。     

Embryological Study on Apomixis in a Sorghum Line SSA-1

The Developmental process of apomictic embryo sac and embryo in a sorghum (Sorghum bicolor (L.) Moench. ) line SSA-1 was observed under light microscope, using the method of conventional paraffin sectioning. The result showed that the apomictic development conforms apospory and diplospory. The uninucleate embryo sac underwent mitotic divisions for three times to form a seven-celled or eight-nuclei mature embryo sac including an egg, two synergids, two polar nuclei and three antipodals. The antipodals divided and multiplicated to form an antipodal mass. Moreover, aposporous multiarchesporial cells and multiple embryo sacs were infrequently observed. Without pollination, the egg divided autonomously to form a typical graminaceous mature embryo. The authors counted the apomictic sections in the whole sections and the result showed that the frequency of apomixis was 42%, indicating the facultative apomictic property in the line SSA-1. The characteristics of apomictic process in the line SSA-1 is also discussed.