ONTOGENY OF THE PALEOZOIC OVULE,CALLOSPERMARION PUSILLUM

The ontogeny of the upper Pennsylvanian age gymnospermous ovule, Callospermarion pusillum, is described from petrifaction specimens collected at the Berryville locality in Illinois. Ovules exhibit a wide range of dimensional and structural features that indicate an extensive developmental sequence. Specimens range from ovules with indistinct zonation of the thin-walled integument to those with thick-walled cells of the sclerotesta. The apex of the fleshy nucellus in some specimens is preserved as a cellular mound, while in others a well-formed cellular pollen chamber is present; still other ovules are characterized by a papery-thin nucellus and pollen chamber wall. The megagametophyte of most specimens is represented by a hollow megaspore membrane that may be restricted to the base of the nucellus, or occupy the entire seed cavity. In a few specimens cellular gametophytes are preserved, and in one ovule archegonia with supposed eggs are also present. Variability in each of the features is compared with ontogenetic changes in comparable structures of living gymnospermous ovules and is correlated with ovule size. A developmental sequence for the fossil ovules is proposed.     

依兰(番荔枝科)雌配子体发育研究

荔枝科是被子植物的基部类群之一,依兰属是番荔枝科较为原始的类群,其有性生殖过程,特别是胚胎发生与发育的研究结果,可以补充被子植物胚胎学原始特征的相关基础资料。该研究利用常规石蜡切片技术,对依兰胚珠、大孢子和雌配子体的发生发育过程进行了观察。结果表明:依兰的胚珠为倒生胚珠、厚珠心、三层珠被,第三层珠被(中间珠被)发生在大孢子母细胞时期,于外珠被与内珠被之间、胚珠合点端两侧发生并隆起;雌配子体为蓼型。此外,依兰的个别胚珠中存在双雌配子体现象,且两个雌配子体均由大孢子母细胞发育而来,大小、形状相近,呈线形排列。该研究结果对于揭示原始被子植物胚胎发育特征具有重要意义。     

The phenotype of Arabidopsis ovule mutants mimics the morphology of primitive seed plants

In seed plants, the ovule is the female reproductive structure, which surrounds and nourishes the gametophyte and embryo. This investigation describes the PRETTY FEW SEEDS2 (PFS2) locus, which regulates ovule patterning. The pfs2 mutant exhibited developmental defects in the maternal integuments and gametophyte. This mutation was inherited as a maternal trait, indicating that gametophyte defects resulted from ovule patterning aberrations. Specifically, the boundary between the chalaza and the nucellus, two regions of the ovule primordia, shifted towards the distal end of pfs2 ovule primordia. Results indicated that the PFS2 locus could: (i) be involved in the development of either the nucellus or the chalaza; or (ii) establish a boundary between these two regions. Examination of genetic interactions of the pfs2 mutation with other well-characterized ovule loci indicates that this locus affects integument morphogenesis. Interestingly, the pfs2 inner no outer and pfs2 strubbelig double mutants had inner integuments that appeared similar to their ancestral precursor. The fossil record indicates that the inner integument evolved by fusion of sterilized sporangia or branches around a central megasporangium. The question of whether the structures observed in these double mutants are homologous or merely analogous to the ancestral precursors of the inner integument is discussed.     

Resolving the systematic and phylogenetic position of isolated ovules: a case study on a new genus from the Permian of China

Isolated ovules occur in many fossil plant assemblages, where they provide important insights into seed‐plant diversity and evolution. However, in many cases, the ovules cannot be attributed to individual groups of seed plants, restricting systematic and evolutionary assessments that can be made from otherwise well‐characterized fossil taxa. In the present paper, we describe a new kind of ovule discovered in tuffaceous sediments from the Permian‐aged Xuanwei Formation in Guizhou Province, China. This ovule has 180° rotational symmetry and an integument comprising a variably thick sarcotesta, a uniformly thick sclerotesta and a uniformly thin endotesta. The nucellus is attached to the integument at least basally and contains a collapsed seed megaspore; a nucellar apex is absent. Both the integument and nucellus are vascularized by paired bundles in the major plane of the ovule; the integumentary bundles are considerably larger than the nucellar bundles and the nucellar bundles emerge from a conical vascular pad. Generation of a three‐dimensional reconstruction based on serial peels revealed the gross morphology and organization of the ovule and highlighted the presence of features consistent with cardiocarpalean‐type ovules (ovule shape, histological features of the integument) and also features more typical of lagenostomalean‐ and trigonocarpalean‐type ovules (large integumentary bundles, presence of nucellar bundles). To assess the affinity and evolutionary significance of the ovule, it has been included in a cladistic matrix of cardiocarpalean‐, lagenostomalean‐ and trigonocarpalean‐type ovules. Results place the ovule within the cardiocarpalean group of ovules known to have been produced by several plant groups, including cordaitean coniferophytes, pteridosperms and Palaeozoic conifers. The cladistic topology supports generic level distinction of the present species, requiring the establishment of Muricosperma guizhouensis Seyfullah & J.Hilton gen. & sp. nov . Lagenostomalean ovules produced by hydrasperman pteridosperms form a basal paraphyletic grade, whereas trigonocarpalean ovules produced by medullosan pteridosperms form a monophyletic group in which Stephanospermum is paraphyletic with respect to Rhynchosperma and Pachytesta. The results also place the Mississippian ovule Mitrospermum bulbosum apart from all of the Pennsylvanian species of Mitrospermum that form a strongly supported clade. Consequently, M. bulbosum is transferred to the new genus Whitaddera Seyfullah & J.Hilton as W. bulbosa (Long) Seyfullah & J.Hilton. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 164 , 84–108.     

Ovule and gametophyte development in Nicotiana glauca Graham (Solanaceae)

Abstract

The development of ovule and megagametophyte is examined in Nicotiana glauca, using light microscopy. The ovules proved unitegmic, tenuinucellate and endothelial as in all the Solanaceae so far studied. The ovule primordia are of the three-zonate type. The integument, which is of dermal origin, is at first two-layered but later produces additional intermediate cells whose origin is not constant. The nucellus, whose initial curvature bears no relation to the origin of the integument, has, like other Solanaceae, a one or two-celled archesporium from which a single meiocyte develops. The gametophyte is confirmed to be bisporic in origin and its development follows the Allium type. Furthermore, the hypostase, which is rare in the family, is observed below the antipodal cells.     

The dyad gene is required for progression through female meiosis in Arabidopsis

In higher plants the gametophyte consists of a gamete in association with a small number of haploid cells, specialized for sexual reproduction. The female gametophyte or embryo sac, is contained within the ovule and develops from a single cell, the megaspore which is formed by meiosis of the megaspore mother cell. The dyad mutant of Arabidopsis, described herein, represents a novel class among female sterile mutants in plants. dyad ovules contain two large cells in place of an embryo sac. The two cells represent the products of a single division of the megaspore mother cell followed by an arrest in further development of the megaspore. We addressed the question of whether the division of the megaspore mother cell in the mutant was meiotic or mitotic by examining the expression of two markers that are normally expressed in the megaspore mother cell during meiosis. Our observations indicate that in dyad, the megaspore mother cell enters but fails to complete meiosis, arresting at the end of meiosis 1 in the majority of ovules. This was corroborated by a direct observation of chromosome segregation during division of the megaspore mother cell, showing that the division is a reductional and not an equational one. In a minority of dyad ovules, the megaspore mother cell does not divide. Pollen development and male fertility in the mutant is normal, as is the rest of the ovule that surrounds the female gametophyte. The embryo sac is also shown to have an influence on the nucellus in wild type. The dyad mutation therefore specifically affects a function that is required in the female germ cell precursor for meiosis. The identification and analysis of mutants specifically affecting female meiosis is an initial step in understanding the molecular mechanisms underlying early events in the pathway of female reproductive development.     

A NOMARSKI INTERFERENCE STUDY OF MEGASPOROGENESIS AND MEGAGAMETOGENESIS IN SMELOWSKIA CALYCINA (CRUCIFERAE)

Ovule development, megasporogenesis and megagametogenesis in an aneuploid population of the arctic-alpine crucifer, Smelowskia calycina, were examined to test for the possibility of apomictic seed production. Whole mounts of ovules cleared in Herr's “4½” clearing fluid were examined using Nomarski differential interference microscopy. The campylotropous ovule was bitegmic, with a micropyle formed by both integuments. The single archesporial cell of a crassinucellar nucellus functioned directly as a megasporocyte, dividing to form a linear tetrad of megaspores. The chalazal megaspore divided to form an 8-nucleate, 7-celled gametophyte of the Polygonum type, having hooked synergids with a well-developed filiform apparatus and polar nuclei that fused prior to fertilization. In the absence of any anomalous development indicative of agamospermy, seed production was assumed to be sexual.     

OVULE ONTOGENY,MEGASPOROGENESIS, AND EARLY GAMETOGENESIS IN TRIFOLIUM REPENS (PAPILIONACEAE)

To aid in understanding of the early events in seed development, surface topography observations with the scanning electron microscope can be coupled with new methods of clearing tissues for light microscopy study. These techniques reveal that two to four ovules begin development along the placental ridge as conduplication of the carpel proceeds in Trifolium repens L. A multicellular archesporium may develop giving rise to several sporogenous cells and ultimately to more than one megasporocyte. However, meiosis is completed in only one megasporocyte to give rise to a single linear tetrad of megaspores. The chalazal megaspore functions in megagametogenesis. Megasporogenesis and megagametogenesis progress as ovule ontogeny proceeds. The outer integument develops more rapidly than the inner and contributes to the final form of the campylotropous ovule. The most dramatic change in ovule form occurs as the tetrad develops and the functional spore enlarges and divides mitotically to produce the two-nucleate megagametophyte. It can be demonstrated that this early gametophyte develops faster than it is allowed to expand in the nucellar mass. This may in part explain why there is gametophyte failure and reduced seed set in clovers.     

The Ovule Structure and Development of Male and Female Gametophytes in Araentotaxus

The ovules of Amentotaxus are 2–2.5 cm in length and about 1.3 cm in width. Aril, which contains about 14—17 secretory canals in transection, is com- posed of parenchyma. The vascular bundles are absent in aril. The integument is com- posed of ceils of outer, middle, and inner regions. The middle region of the integument is about 10 cells in thickness. The Integuments are basally parenchymatous before pollination and then the cells become elongating and thickenning in middle region after pollination. Finally the micropylar canal is forced to be closed. In addition, there are 8–14 vascular bundles with centripetal xylem arranged in a ring in integument. One of the most remarkable feature of the ovule is that there is a pollen chamber formed at the top of nucellus before pollination. At this time because epidermis of nucellus does not disintegrate, the pollen chamber looks like conical in longitudinal section. After pollination the pollen chamber was becoming closed due to elongation and thickenning of cells in the middle region of integument. At the base of ovule there are 4–5 pairs of bracts which contain a larger secretory cavity and a centripetal xylem in a vascular bundle. It is interesting that wax layer of 30–40 μm thick is covered on the surface of integument, aril and bract. Usually 3–4 microsporangia which are hypodermal in origin, occur in abaxial side of a microsporophyll. In some cases the tapetum is partly enlarging and extruding into the developing tetrahedral tetrads. The mature pollen comprises an antheridial initial and a tube cell. About 20 pollen grains may germinate in the same ovule. The megaspore divide successively 8 times to produce 256 nuclei and then cell wall formation takes place. The female gametophyte is about 830–908 μm in length and 500 μm in width. The archegonia are single, terminal, and 6–7 in number. The mature archgonium, with ventral canal nucleus, is about 430 μm in length and 80–108 μm in width. The female gametophyte is often growing against the upper part of the nuceilus and makes the cells of the latter gradually to be disintegrated. The ovule construction of Amentotaxus is in some degree similar to that of Ginkgo in having a comparatively well developed pollen chamber. The mature pollen of Amentotaxus, which is similar to that of Cephalotaxus is composed of 2-cells. In sum, Amentotaxus perhaps is the most primitive genus in Taxaceae and it is closely related to Cephalotaxus.     

Angiosperm ovules: diversity, development, evolution

Background

Ovules as developmental precursors of seeds are organs of central importance in angiosperm flowers and can be traced back in evolution to the earliest seed plants. Angiosperm ovules are diverse in their position in the ovary, nucellus thickness, number and thickness of integuments, degree and direction of curvature, and histological differentiations. There is a large body of literature on this diversity, and various views on its evolution have been proposed over the course of time. Most recently evo–devo studies have been concentrated on molecular developmental genetics in ovules of model plants.

Scope

The present review provides a synthetic treatment of several aspects of the sporophytic part of ovule diversity, development and evolution, based on extensive research on the vast original literature and on experience from my own comparative studies in a broad range of angiosperm clades.

Conclusions

In angiosperms the presence of an outer integument appears to be instrumental for ovule curvature, as indicated from studies on ovule diversity through the major clades of angiosperms, molecular developmental genetics in model species, abnormal ovules in a broad range of angiosperms, and comparison with gymnosperms with curved ovules. Lobation of integuments is not an atavism indicating evolution from telomes, but simply a morphogenetic constraint from the necessity of closure of the micropyle. Ovule shape is partly dependent on locule architecture, which is especially indicated by the occurrence of orthotropous ovules. Some ovule features are even more conservative than earlier assumed and thus of special interest in angiosperm macrosystematics.     

濒危植物马蹄参的大孢子发生和雌配子体发育

为探讨马蹄参(Diplopanax stachyanthus Hand.-Mazz.)濒危机制与雌性生殖发育的关系,采用石蜡切片法观察马蹄参大孢子形成和雌配子体的发育过程。结果表明,马蹄参雌蕊单心皮,子房下位,1室,1枚胚珠。胚珠横生于短片状胎座上,具单珠被,厚珠心。单孢原细胞自珠心1~2层表皮细胞处分化;大孢子四分体为直线形。成熟胚囊中,2个极核在受精前融合为次生核,3个反足细胞不发达,较早退化;二核胚囊时期出现二核分裂不均且较小核退化消失的异常发育现象。因此,马蹄参雌配子体发育过程中出现异常现象是造成其结实率低的主要原因。     

Comparative embryology of Bambusa tulda Roxb. and Thyrsostachys siamensis Gamble (Poaceae: Bambuseae)

Bambusa tulda and Thyrsostachys siamensis resemble each other in having an obovate ovary which is hairy and thickened along the apex, a pseudo-crassinucellate ovule with a wide region of attachment, poorly-developed and ephemeral outer integument, an inner integument which fails to grow beyond the nucellus, 'Polygonum' type of embryo sac ontogeny, parallel orientation of embryo sac to the long axis of the ovule, multiple antipodals which retain apical position in the embryo sac even during post-fertilization phase of development, an ephemeral nucellus, relatively small bambusoid embryos, and many-layered and apically thickened pericarp. However, they differ from each other in their gynoecial structure, the extent of the development of the outer integument, organization of megaspore tetrads and development-stage-related behaviour of the inner integument in the fertilized ovules. These taxa also differ from other members of the subfamily Bambusoideae in the structure of the mature ovule, endosperm and pericarp.     

矮沙冬青雌配子体及胚胎发育研究

矮沙冬青子房单心皮1室,边缘胎座,弯生胚珠,胚珠具双珠被、厚珠心。大孢子孢原细胞发生于珠心表皮下,大孢子母细胞减数分裂形成直线排列的四分体,合点端大孢子具功能,并按蓼型胚囊发育,雌配子体成熟于4月中旬。双受精后,胚乳发育为核型。在矮沙冬青大孢子发生、雌配子体和胚胎发育过程中未发现异常现象,因此认为矮沙冬青濒危不存在雌性生殖结构与发育过程异常的内在因素。     

Latisemenia longshania,gen. et sp. nov., a new Late Devonian seed plant from China

The earliest known ovules in the Late Devonian (Famennian) are borne terminally on fertile branches and are typically enclosed in a cupule. Among these ovules are some that have terete integumentary lobes with little or no fusion. Here, we report a new taxon, Latisemenia longshania, from the Famennian of South China, which bears cupulate ovules that are terminal as well as opposite on the fertile axis. Each ovule has four broad integumentary lobes, which are extensively fused to each other and also to the nucellus. The cupule is uniovulate, and the five flattened cupule segments of each terminal ovule are elongate cuneate and shorter than the ovule. Associated but not attached pinnules are laminate and Sphenopteris-like, with an entire or lobate margin. Latisemenia is the earliest known plant with ovules borne on the side of the fertile axis and may foreshadow the diverse ovule arrangements found among younger seed plant lineages that emerge in the Carboniferous. Following the telome theory, Latisemenia demonstrates derived features in both ovules and cupules, and the shape and fusion of integumentary lobes suggest effective pollination and protection to the nucellus. Along with other recent discoveries from China, Latisemenia extends the palaeogeographic range of the earliest seed plants.     

MITROSPERMUM VINCULUM SP. NOV., A CARDIOCARPALEAN OVULE FROM THE UPPER PENNSYLVANIAN OF OHIO

Numerous anatomically preserved ovules assignable to the genus Mitrospermum have been discovered in Upper Pennsylvanian sediments of Eastern Ohio. Although basically similar to Mitrospermum compressum, the newly discovered specimens exhibit several consistent differences. Ovules are strongly platyspermic, up to 4.2 mm long, 4.0 mm wide, and 0.6 mm thick. In the minor plane, ovules are broadest at the base and taper toward the micropyle. The integument exhibits three topographic regions: endotesta, sclerotesta, and sarcotesta. The sarcotesta is extremely broad in the major plane, where it forms two membranous wings. A single terete vascular bundle enters the base of the ovule, traverses the integument, and divides to form two integumentary bundles and a conspicuous nucellar platform. Integumentary bundles extend toward the tip of the ovule at the margin of the sarcotesta and sclerotesta. A pollen chamber with a prominent nucellar beak is delimited at the tip of the nucellus. Consistent differences in vascularization, size, nature of the seed base, features of the pollen chamber, and the Late Pennsylvanian age demonstrate that the specimens represent a distinct species. The discovery of these ovules extends the stratigraphic range of Mitrospermum to the Upper Pennsylvanian of Ohio.     

小盐芥大孢子发生和雌配子体发育

本论文研究了小盐芥(Thellungiella halophila)大孢子发生和雌配子体发育过程及该阶段与花蕾、花、果实外部形态的相关性。结果如下: 小盐芥雌蕊由2心皮组成, 侧膜胎座, 每室胚珠多数, 弯生, 双珠被, 薄珠心。孢原细胞位于珠心表皮之下, 直接起大孢子母细胞的功能。大孢子四分体线形排列, 合点端大孢子为功能大孢子, 胚囊发育为蓼型。     

小叶兜兰的果实生长和雌配子体发育

为了解濒危兰科植物小叶兜兰(Paphiopedilum barbigerum Tang et Wang)胚珠和雌配子体的发育过程,采用常规石蜡切片技术对其果实的生长动态进行了研究。结果表明,授粉后60~75 d的蒴果内种子数量迅速增加,到授粉后120 d时种子充满整个蒴果。授粉后40 d的胎座上分化形成多数由1层表皮细胞包被1列细胞的胚珠原基;授粉后60 d时位于胎座指状结构末端处紧靠表皮细胞下方的孢原细胞分化为大孢子母细胞。之后,大孢子母细胞经过减数分裂和有丝分裂最终形成成熟胚囊;授粉后135 d胚囊发育成熟,附着在胎座上的种子个体分化明显。小叶兜兰胚囊的发育类型为双孢子葱型,胚珠为倒生胚珠,薄珠心,单珠被,成熟胚囊为8核。这为小叶兜兰的生殖生物学及繁殖体系的建立提供理论依据。     

小盐芥大孢子发生和雌配子体发育

本论文研究了小盐芥（Thellungiella halophila）大孢子发生和雌配子体发育过程及该阶段与花蕾、花、果实外部形态的相关性。结果如下：小盐芥雌蕊由2心皮组成,侧膜胎座,每室胚珠多数,弯生,双珠被,薄珠心。孢原细胞位于珠心表皮之下,直接起大孢子母细胞的功能。大孢子四分体线形排列,合点端大孢子为功能大孢子,胚囊发育为蓼型。     

罗汉果大孢子发生及雌配子体发育与花部形态、胚珠变化的关系

为弄清罗汉果(Siraitia grosvenorii)大孢子发生、雌配子体发育过程与花部形态特征、胚珠的关系,运用石蜡切片法对罗汉果子房进行了显微观察。结果表明,罗汉果的胚珠倒生,双珠被,厚珠心,大孢子四分体呈线型排列,合点端一个大孢子分化为功能大孢子,成熟胚囊为蓼型。花蕾形态、胚珠变化与大孢子发生、雌配子体的发育时期具有一定相关性,当子房长度为7.0 mm≤L<9.0 mm,珠心呈椭圆形时,约有45.83%的大孢子母细胞处于减数分裂时期。因此,依据罗汉果花部形态可有效确定大孢子发生与雌配子体发育的时期。     

Temporal and Spatial Expression of MADS Box Genes, FBP7 and FBP11, During Initiation and Early Development of Ovules in Wild Type and Mutant Petunia hybrida

Abstract: The temporal and spatial distribution of the Petunia Floral Binding Proteins 7 and 11 (FBP7/11) were determined immunocytochemically during ovule initiation and development. In wild type plants, FBP7/11 were first detected in the placenta before ovule primordia were formed. At ovule primordium stage, FBP7/11 levels increased in the placenta and appeared in ovule primordia at the sites where integument primordia developed. At the megagametogenesis stage, FBP7/11 were present at high levels in the placenta, funicle and integument, but not in the nucellus or gametophyte. Transgenics with cosuppression of FBP7/11 formed normal ovule primordia on the placenta from which both normal ovules and carpel-like structures developed. The amount of FBP7/11 was low in the ovules and undetectable in the carpel-like structures. Plants with ectopic expression of FBP7/11 developed normal ovules on the placenta and, in addition, ovule- and carpel-like structures on sepals. Placental and sepal ovules showed the same labeling pattern as observed in wild type ovules. FBP7/11 levels were, however, low or undetectable in the carpel-like structures. The results indicate that FBP7/11 only have indirect roles in ovule primordium initiation. However, at least small quantities are needed for proper ovule differentiation. Thus, the amount of FBP7/11 is related to the type of development after primordium formation, i.e., towards the formation of real ovules or carpel-like structures.