几种具有无融合生殖特性的植物多胚和多苗现象的观察

报道６种具无融合生殖特性的植物种子的胚数和萌发实生苗数的观察结果。金桔、密桔、和花椒具珠心胚,含多胚种子频率分别为９７．５％、１００％和５４．００％,多胚种子的胚数范围分别为２￣４９、３￣５４和２￣６。草地早熟禾、滨草和湖北海棠具非减数配子体无融合生殖特性,含多胚种子频率依次为４３．２５％、８．１１％和３７．５０％,前两种的多胚种子中胚数范围为２￣３,后为２￣１５。密桔、草地早熟禾和湖北海棠种子     

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。     

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     

草地早熟禾无融合生殖现象的研究

本文应用石蜡切片和子房整体染色透明法研究了草地早熟禾(Poa pratensisL.)的无融合生殖现象,结果表明,草地早熟禾的胚胎发育分为两种类型。一种是由孤雌生殖形成胚,即卵细胞分裂较早形成幼胚,少数胚囊中的一个或两个助细胞也分裂较早,同时形成幼胚,但两个极核并不融合,三个反足细胞明显增大。由孤雌生殖形成胚的约占观察总数的66%,其中3%为双胚,仅一个是三胚。在成熟种子中也有两个成熟胚并生的现象。另一种是形成胚和胚乳,约占34%。34%中形成种子有两种可能的途径,一是通过有性生殖形成种子,二是由孤雌生殖形成的胚和极核受精形成的胚乳共同组成的无融合生殖种子。本研究还首次观察到早熟禾完整的双胚囊结构。     

草地早熟禾胚胎学研究 Ⅲ．多胚囊及多胚现象

报道了草地早熟禾中多胚囊的起源、发育和结构。在１个胚珠中,大孢子母细胞周围可以有一到多个起源于珠心细胞的胚囊原始细胞,并可以发育成为多胚囊,其中具有两个胚囊的可以发育成为成熟胚囊。起源于珠心的体细胞无孢子生殖胚囊的发育属于山柳菊型。两个成熟胚囊中,都可以形成胚和胚乳,因而形成了具假多胚的种子。位于中部的胚来源于珠心还囊,属于无融合生殖形成的胚。两个以上的多胚囊不能形成成熟胚囊。     

多胚水稻胚位与苗位的观察研究

多胚水稻是进行水稻无融合生殖研究的重要材料,通过对4个多胚水稻品系种子胚位及萌发后的苗位观察发现这些品系中存在着多种胚位及苗位类型,其中以双-13的类型最为丰富。认为不定胚在与合子胚共存的情况下难以萌发表达,而只有当合子胚败育时才有可能得以萌发,并且认为反向单苗,侧向单苗及倒序单苗可能起源于不定胚,因而可能成为进行水稻无融合生殖筛选的重要标志。     

草地早熟禾的多胚现象

草地早熟禾的多胚现象母锡金,王伏雄,梁铁兵（中国科学院植物研究所,北京１０００４４）草地早熟禾（Ｐｏａｐｒａｔｅｎｓｉｓ）无融合生殖的研究历时半个多世纪,近年来又有新的进展（ＢａｓｈａｗａｎｄＨａｎｎａ,１９９０）。为了同水稻“无融合多胚苗”进行比较...     

提高水稻双胚苗和不定胚频率的研究

以水稻双胚苗中提高不定胚的发生频率是研究利用无融合生殖固定F_1代杂种优势的主要途径之一。通过特殊的选育方法,已将原有双胚苗品系的双苗率由30％提高到60％左右。由子不定胚每双苗的发生率有着一定的内在联系,随着双苗率的提高,内含不定胚的频率也有相应提高的趋势。又据细胞学观察,认为不定胚发生在远离珠孔端的部位,故在种子中部的萌发苗可能是不定胚产生的种子形态学特征。今年,我们采用特定苗位作标记的筛选方法,选取种子正中部萌发苗并结合对其它非正中苗的人工切除,使水稻的不定胚频率由原来的2.6-5.1％提高到21.2％。水稻不定胚频率的大幅度提高,为无融合生殖在杂交水稻育种中的研究应用开拓了诱人的前景。     

青藏高原东部植物种子萌发能力与其在群落中相对多度的关系

不同演替阶段的群落中植物幼苗建植特性以及幼苗建植时所处的环境都会有所不同.这些差异可能会导致种子萌发率与物种多度之间的关系随着群落的演替阶段而发生改变.目前关于不同类型群落中以及同一类型群落的不同演替阶段物种多度和种子萌发能力之间的关系的研究较少.本研究以全面收集的青藏高原东缘3种常见次生植被(鼢鼠土丘群落、人工草地群落和弃耕地群落)中植物种子为基础,沿着群落演替梯度来探明种子萌发和物种多度之间的关系.通过对结果的线性回归分析发现:在鼢鼠土丘群落和人工草地群落中,种子萌发率和物种多度之间无显著相关性;而在弃耕地群落中,种子萌发率和物种多度之间的关系有随着群落的演替呈现由正相关向负相关转变的趋势.     

几个芒果品种的胚性及多胚苗遗传分析

通过目测,筛选出芒果多胚种子,观察其多胚形态;将多胚种子培育成多胚苗,观察其生长发育状况;取多胚苗和母树叶片进行体细胞染色体数目鉴定及同工酶的分析.结果表明,多胚类群的品种中有单胚现象出现,土芒、吕宋、象牙单胚出现的频率分别为1.04%、3.85%和5.44%;土芒和象牙胚数出现的范围分别为2-6和2-8.多胚种子子叶形态和胚轴的着生位置各异.种子多胚的萌发率、成活率及多胚苗的生长状况均与多胚种子的胚数、子叶发育大小等因素相关.多胚苗体细胞染色体数目是2n=40,未发现染色体的数目变异.POD同工酶分析表明,同一种子的多胚苗间酶谱上存在着差异.     

Polyembryony in Citrus. Accumulation of seed storage proteins in seeds and in embryos cultured in vitro.

Citrus exhibits polyembryonic seed development, an apomictic process in which many maternally derived embryos arise from the nucellus surrounding the developing zygotic embryo. Citrus seed storage proteins were used as markers to compare embryogenesis in developing seeds and somatic embryogenesis in vitro. The salt-soluble, globulin protein fraction (designated citrin) was purified from Citrus sinensis cv Valencia seeds. Citrins separated into two subunits averaging 22 and 33 kD under denaturing sodium dodecyl sulfate-polyacrylamide gel electrophoresis. A cDNA clone was isolated representing a citrin gene expressed in seeds when the majority of embryos were at the early globular stage of embryo development. The predicted protein sequence was most related to the globulin seed storage proteins of pumpkin and cotton. Accumulation of 33-kD polypeptides was first detected in polyembryonic Valencia seeds when the majority of embryos were at the globular stage of development. Somatic Citrus embryos cultured in vivo were observed to initiate 33-kD polypeptide accumulation later in embryo development but accumulated these peptides at only 10 to 20% of the level observed in polyembryonic seeds. Therefore, factors within the seed environment must influence the higher quantitative levels of citrin accumulation in nucellar embryos developing in vivo, even though nucellar embryos, like somatic embryos, are not derived from fertilization events.     

Polyembryony and apomixis in Eriotheca pubescens (Malvaceae - Bombacoideae)

Apomixis and adventitious polyembryony have been reported for several species of Bombacoideae, including Eriotheca pubescens, a tree species of the Neotropical savanna (Cerrado) areas in Brazil. However, the origin of polyembryonic seeds and their importance for the reproduction of the species remained to be shown. Here, we analyzed the early embryology of this species to establish the apomictic origin of extranumerary embryos. We also observed the geographic distribution of polyembryony in E. pubescens, and tested if apomixis was related to the source of pollen (self or cross) and population density. Moreover, we tested if polyembryonic apomictic embryos would develop normally into seedlings. In the observed seed primordia, after a relatively long quiescent period, the zygote developed into a sexual embryo concurrently with adventitious apomictic embryos which developed from nucellus cells. Adventitious embryos develop faster than sexual ones and are morphologically similar, so that 44 days after anthesis it was virtually impossible to distinguish and trace the fate of the sexual embryo. Polyembryony is widely distributed in populations some 400 km distant, and only one strictly monoembryonic individual was observed during the study. The number of embryos per seed varied between fruits and individuals but was significantly higher in seeds from cross-pollinations than from selfs, although fruit and seed set after crosses were much lower than after selfs. Embryo development into seedlings depended on their weight at germination, but polyembryonic seeds germinated and produced up to seven seedlings per seed in greenhouse conditions. Adventitious embryony and apomictic seedlings would explain the mostly clonal populations suggested by molecular studies.     

AFLP markers closely linked to a major gene essential for nucellar embryony (apomixis<Emphasis Type="Italic">)</Emphasis> in <Emphasis Type="Italic">Citrus maxima</Emphasis> × <Emphasis Type="Italic">Poncirus trifoliata</Emphasis>

Some citrus varieties express a form of apomixis termed nucellar embryony in which the adventive embryos develop from nucellus tissue surrounding the embryo sac. This trait results in many seeds containing multiple embryos (polyembryony). Inheritance of the frequency of polyembryony was studied in 88 progeny from a cross of Citrus maxima (monoembryonic) × Poncirus trifoliata (polyembryonic). The frequency of polyembryonic seed produced by each progeny was determined by scoring 100–500 seeds for the number of seedlings to emerge from each seed. Two groups of eight individuals from each extreme of the population were chosen for bulked segregant analysis with amplified fragment length polymorphism markers amplified with 256 primer combinations. Candidate markers identified in the bulks as linked to the trait were tested on the 32 individuals used to create the bulks and then on the remaining plants in the population. Five candidate markers tightly linked to polyembryony in P. trifoliata were identified. Specific marker alleles were present in nearly all progeny that produced polyembryonic seed, and alternate alleles were present in nearly all progeny that produced only monoembryonic seed. The region defined by these markers very likely contains a gene that is essential for the production of polyembryonic seeds by apomixis, but also shows segregation distortion. The proportion of polyembryonic seeds varied widely among the hybrid progeny, probably due to other genes. Scoring 119 progeny of a P. trifoliata selfed population for the closely linked markers and the proportion of polyembryonic seeds confirmed close linkage between these markers and polyembryony.     

Assessment of polyembryony in lemon: rescue and in vitro culture of immature embryos

The 27 lemon cultivars analysed could be considered slightly or moderately polyembryonic, with 25 to 43% of seeds being polyembryonic and from 1.3 to 1.6 embryos per seed. On this basis, it is necessary to rescue zygotic embryos at an immature stage. Rescue and in vitro embryo development have been studied in two Citrus limon polyembryonic cultivars. Sucrose (50 and 70 g/l) was combined with Murashige and Skoog and Gamborg’s B5 media and tested for optimal growth response. An important effect of genotype was observed: embryos from cultivar ‘Eureka’ had greater survival, germination percentage, and radical development. While the sucrose concentration in the medium did not have an effect on germination, the medium affected the embryo survival and root development of the seedlings, Gamborg’s B5 medium giving the best results. The ability to form plants in vitro was affected by an increase of embryo developmental stage. The germination and seedling height were greater with embryos of seeds collected 135–150 days after anthesis.     

Sexual and apomictic development in Hieracium

 Most members of the genus Hieracium are apomictic and set seed without fertilization, but sexual forms also exist. A cytological study was conducted on an apomictic accession of H. aurantiacum (A3.4) and also H. piloselloides (D3) to precisely define the cellular basis for apomixis. The apomictic events were compared with the sexual events in a self-incompatible isolate of H. pilosella (P4). All plants were maintained as vegetatively propagated lines each derived from a single plant. Sexual P4 exhibited characteristic events of polygonum-type embryo sac formation, showed no latent apomitic tendencies, and depended upon fertilization to set seed. In contrast, D3 and A3.4 were autonomous aposporous apomicts, forming both embryo and endosperm spontaneously inside an unreduced embryo sac. The two apomicts exhibited distinct mechanisms, but variation was also observed within each apomictic line. Seeds from apomicts often contained more than one embryo. A degree of developmental instability was also observed amongst germinated seedlings and included variation in meristem and cotyledon number, altered phyllotaxis, callus formation, and seedling fusion. In most cases abnormal seedlings developed into normal plants. Such phenomena were not observed following germination of hybrid seeds derived from crosses between sexual P4 and the apomictic plants. The three plants can now be used in inheritance studies and also to investigate the molecular mechanisms controlling apomixis. Received: 11 February 1998 / Revision accepted: 23 July 1998     

Embryological Observation on the Apomictic Rice-ApⅢ (Shuang 13)

Since 1988s, the phenomenon of multiple seedlings and multiple embryoes of apomictic rice, Ap Ⅲ (Shuang 13) has been interestingly concerned. Embryological investigation of Ap Ⅲ has been carried out with the whole dissection and conventional sectioning technique. The main conclusions are as follows: 1.8. 9 % of 5000 matured caryopsises, contained one embryo in a caryopsis, from which single seedling arose, while 8.9 % twin embryoes and 1.2 % triplet embryoes of which developed two seedlings and triple seedlings respectively; 2. In over 700 young caryopsises, 90.0%～95.0% contained only one embryo, 5.0%～7.0% twin embryoes, except a few of the ovules with absorptive embryo sac or embryo and endosperm. Triplet embryoes caryopsis could not be traced further due to their low frequency; 3. Authors' materials showed that all the single, twin and triplet embryoes were situated at the micropylar end in one embryo sac of young ovule and no adventitious embryo was seen outside the embryo sac of ovules. From the aforementioned results, it can be infered that albeit single embryo is derived from the egg cell yet it could be possible twin or triplet embryocs may be developed from the egg cell and one or two of the synergids either or not through fertilization.     

Sexual,apomictic and mixed populations in <Emphasis Type="Italic">Handroanthus ochraceus</Emphasis> (Bignoniaceae) polyploid complex

Some tropical Bignoniaceae form sporophytic apomictic polyploid complexes are similar to better studied temperate plants. Handroanthus ochraceus is a widely distributed Neotropical savanna tree with both monoembryonic/self-sterile, and polyembryonic/apomictic and self-fertile populations, but lacking chromosome number and morphological comparative studies. We tested if monoembryonic/non-apomictic and polyembryonic/apomictic populations differed in ploidy and morphological features, as a basis to understand evolution and biogeography of these plants. Chromosome number and embryo number per seed were investigated, and uni- and multivariate analyses of flower and leaf morphology were done for five populations of H. ochraceus. We found two pure monoembryonic diploid (2n?=?40), and one pure polyembryonic (62–94% of polyembryonic seeds) tetraploid population (2n?=?80). One of the diploid populations presented only one individual with 3.2% polyembryonic seeds and was considered a non-apomictic population. Another population showed predominantly polyembryonic (27–66% of polyembryonic seeds) tetraploid individuals, but one diploid individual with 2% of polyembryonic seeds, and was considered a mixed apomictic and non-apomictic population. Morphological analyses confirmed breeding system clusters, and that stigma width, as well as pollen grain area, was consistently larger in polyembryonic populations. Polyploid plants showed larger cells, as well as larger organs and other distinctive features, which will be useful to identify apomictic populations and to future taxonomic discussions. The species can be considered an agamic complex with apomixis associated with neopolyploidy. This trend is also found in other tropical sporophytic apomictics, contrasting with the usual reports linking diploidy or paleopolyploidy to this kind of apomictics.     

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.