COMPARATIVE MORPHOLOGY OF THE CARPEL IN THE ROSACEAE. II. PRUNOIDEAE: MADDENIA,PYGEUM; OSMARONIA

A survey of species of the prunoid genera, Maddenia and Pygeum, and of the genus Osmaronia has been made. The ovules of all are pendent, campylotropous, and epitropic. In the prunoids, the ovular supply is intimately connected with a central vascular plexus in the base of the carpel; that plexus is absent from Osmaronia. The prunoid carpels are marked by an extensive degree of fusion among the ovular and wing bundles, by fusion of the sutural margins, by fusion of the 2 integuments of the ovule to a single massive one, and by the presence of 3 or 5 well-developed bundles in the base. The carpel of Osmaronia also has a strongly fused bipartite ovular supply, separate bundles of which, however, become very much attenuated before reaching the funiculus; it has independent ovular and wing bundles, completely separate carpellary margins, 2 clearly separate integuments in the ovule, and 6 distinctive bundles in the carpel base. At the funiculus, the wing bundle of Osmaronia is connected with the adjoining weak ovular bundle by a well-developed vascular branch. Various particularities in the morphology of Osmaronia lend support to its segregation into a unique tribe, the Osmaronieae of Rydberg.     

A structural investigation of the ovule in sugar beet, Beta vulgaris: Integuments and micropyle

The micropyle and the integuments of sugar beet (Beta vulgaris) ovules have been investigated by light and electron microscopy during differentiation and maturation of the ovule. The micropyle itself is formed by the inner integument which is surrounded by the outer integument at its base. The micropyle containts a fibrillar PAS+ substance and is often covered by a thin sheet or hymen. Both integuments are cuticle-covered thin sheets, each 2-few cell layers in thickness. In the outer integument an increase in starch accumulation occurs during ovule maturation and probably functions as nutrient storage for embryo development. The inner epidermis of the inner integument differentiates as the most conspicuous cell layer of the beet ovule. During growth and maturation of the ovule a system of small perinuclear vacuoles containing dense material increases steadily in these cells. At maturity this system fills up more than half of each cell and very dense material has accumulated in each vacuole. This vacuole content is highly refractive and contains tannins and/or polyphenols.     

Interactions among Genes Regulating Ovule Development in Arabidopsis Thaliana

The INNER NO OUTER (INO) and AINTEGUMENTA (ANT) genes are essential for ovule integument development in Arabidopsis thaliana. Ovules of ino mutants initiate two integument primordia, but the outer integument primordium forms on the opposite side of the ovule from the normal location and undergoes no further development. The inner integument appears to develop normally, resulting in erect, unitegmic ovules that resemble those of gymnosperms. ino plants are partially fertile and produce seeds with altered surface topography, demonstrating a lineage dependence in development of the testa. ant mutations affect initiation of both integuments. The strongest of five new ant alleles we have isolated produces ovules that lack integuments and fail to complete megasporogenesis. ant mutations also affect flower development, resulting in narrow petals and the absence of one or both lateral stamens. Characterization of double mutants between ant, ino and other mutations affecting ovule development has enabled the construction of a model for genetic control of ovule development. This model proposes parallel independent regulatory pathways for a number of aspects of this process, a dependence on the presence of an inner integument for development of the embryo sac, and the existence of additional genes regulating ovule development.     

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.     

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.     

非洲狼尾草胚珠(着重胚珠附器)发育的特点

用光学显微镜对非洲狼尾草胚珠发育做了进一步观察。结果表明有如下几个特点：（１）珠孔区域的珠心细胞发生特化,膨大１伸长并进入珠孔,形成胚珠附器。（２）珠心表皮平周分裂产生周缘珠心组织。（３）珠孔由内珠被和腹侧外珠被构成。     

Ovule morphogenesis in Ranunculaceae and its systematic significance

BACKGROUND AND AIMS: Ranunculaceae has a prominent phylogenetic position in Ranunculales which appears at the base of eudicots. The aims of the present paper are to reveal the features of ovule morphogenesis in different taxa and gain a better understanding of the systematics of Ranunculaceae. METHODS: Flowers of 17 species from three subfamilies, nine tribes and 16 genera of Ranunculaceae, at successive developmental stages, were collected in the wild and studied with a scanning electron microscope. KEY RESULTS: The integuments in the unitegmic ovules in Helleborus, Ranunculus and Oxygraphis, as well as the inner integuments in the bitegmic genera, initiate annularly and eventually become cup-shaped. However, the integuments in the unitegmic ovules in Anemone and Clematis, as well as the outer integuments in the bitegmic genera, arise semi-annularly and eventually become hood-shaped. Different kinds of appendages appear on the ovules during development. In Coptis of subfamily Coptidoideae, a wrap-shaped appendage arises outside the ovule and envelopes the ovule entirely. In the genera of subfamily Thalictroideae and tribe Anemoneae of subfamily Ranunculoideae, appendages appear on the placenta, the funicle or both. In tribe Helleboreae of subfamily Ranunculoideae, an alary appendage is initiated where the integument and the funicle join and becomes hood-shaped. CONCLUSIONS: Ovule morphogenesis characteristics are significant in classification at the levels of subfamilies and tribes. The initiation patterns of the integuments and the development of appendages show diversity in Ranunculaceae. The present observations suggest that the bitegmic, hood-shaped outer integument and endostomic micropyle are primitive while the unitegmic, cupular-shaped outer integument and bistomic micropyle are derivative.     

BELL1 and AGAMOUS genes promote ovule identity in Arabidopsis thaliana

Molecular and genetic analyses have demonstrated that the Arabidopsis thaliana gene BELL1 (BEL1) is required for proper morphogenesis of the ovule integuments. Several lines of evidence suggest that BEL1 may act, at least in part, to repress the function of the organ identity gene AGAMOUS (AG) during ovule development. To study the relative roles of BEL1 and AG, plants homozygous for ag, bel1 or both were constructed in an ap2 mutant background where ovules form even in the absence of AG function. The loss of either BEL1 or AG led to a decrease in the number of mature ovules, accompanied by an increase in primordial outgrowths. These data suggest that BEL1 and AG gene products act early in ovule development in a partially redundant manner to direct ovule identity. Development of the abnormal integuments characteristic of the Bel1- mutant phenotype was found to be dependent on AG function. Finally, BEL1 appears to be required for embryo sac development independent of both other aspects of ovule morphogenesis and AG function. This study therefore suggests that both BEL1 and AG are required for several distinct aspects of ovule morphogenesis.     

EMBRYOLOGICAL STUDIES IN THE FAMILY SAXIFRAGACEAE (S.L.). I. DEVELOPMENT OF THE OVULE AND EMBRYO SAC IN SAXIFRAGA FORTUNEI VAR. PARTITA (MAKINO) NAKAI

The development of the ovule, megaspore and megagametophyte in Saxifraga fortunei var. partita (Makino) Nakai was observed. The ovule is anatropous, bitegmic, and crassinucellate. Both integuments originate from the epidermis. The archesporium is considered to be multicellular. The primary sporogenous cell functions as the megaspore mother cell which forms a T-shaped tetrad. The chalazal member of the megaspore tetrad is functional and develops into a Polygonum-type embryo sac. In the pyriform synergids the filiform apparatus is observed, but any hook or indentations could not be recognized. The antipodal cells are detectable until the Helobial endosperm undergoes several nuclear divisions. Secondary multiplication of the nuclei or the cells of the antipodals could not be observed.     

重楼属两种植物种子及其附属结构的发育

重楼属两种植物(五指莲Paris axialis和滇重楼Paris polyphylla var.yunnanensis)种子发育的过程基本一致。双受精发生于授粉后10—15天。胚乳为沼生目型。种子发育延续的时间约为150—170天。胚胎发育终止于球形或稍有分化的阶段。种子具二层种皮。 二种重楼种子成熟时的外部形态显著不同。五指莲Paris axialis的种子呈浅棕黄色,长椭圆形,部分为绿白色海绵质假种皮所包裹。假种皮由珠柄发育而来,呈楔形。滇重楼Parispolyphylla var.yunnanensis的种子鲜红色,不规则圆形,外种皮肉质多浆。无假种皮。珠柄橙黄色,短而纤细。     

Genetic and molecular interactions between BELL1 and MADS box factors support ovule development in Arabidopsis

In Arabidopsis thaliana and many other plant species, ovules arise from carpel tissue as new meristematic formations. Cell fate in proliferating ovule primordia is specified by particular ovule identity factors, such as the homeodomain factor BELL1 (BEL1) and MADS box family members SEEDSTICK (STK), SHATTERPROOF1 (SHP1), SHP2, and AGAMOUS. Both in the bel1 mutant and the stk shp1 shp2 triple mutant, integuments are transformed into carpelloid structures. Combining these mutants in a bel1 stk shp1 shp2 quadruple mutant, we showed that the bel1 phenotype is significantly enhanced. We also demonstrate that ovule differentiation requires the regulation of the stem cell maintenance gene WUSCHEL, repression of which is predominantly maintained by BEL1 during ovule development. Based on yeast three-hybrid assays and genetic data, we show that BEL1 interacts with the ovule identity MADS box factors when they dimerize with SEPALLATA proteins. We propose a model for ovule development that explains how the balance between carpel identity activity and ovule identity activity is established by a MADS box homeodomain protein complex.     

Expression-based discovery of candidate ovule development regulators through transcriptional profiling of ovule mutants

Background  

Arabidopsis ovules comprise four morphologically distinct parts: the nucellus, which contains the embryo sac, two integuments that become the seed coat, and the funiculus that anchors the ovule within the carpel. Analysis of developmental mutants has shown that ovule morphogenesis relies on tightly regulated genetic interactions that can serve as a model for developmental regulation. Redundancy, pleiotropic effects and subtle phenotypes may preclude identification of mutants affecting some processes in screens for phenotypic changes. Expression-based gene discovery can be used access such obscured genes.     

Megasporogenesis,megagametogenesis, and embryogenesis in Dendrobium nobile (Orchidaceae)

The orchid reproductive strategy, including the formation of numerous tiny seeds, is achieved by the elimination of some stages in the early plant embryogenesis. In this study, we documented in detail the formation of the maternal tissues (the nucellus and integuments), the structures of female gametophyte (megaspores, chalazal nuclei, synergids, polar nuclei), and embryonic structures in Dendrobium nobile. The ovary is unilocular, and the ovule primordia are formed in the placenta before the pollination. The ovule is medionucellate: the two-cell postament and two rows of nucellar cells persist until the death of the inner integument. A monosporic eight-nucleated embryo sac is developed. After the fertilization, the most common central cell nucleus consisted of two joined but not fused polar nuclei. The embryogenesis of D. nobile is similar to the Caryophyllad-type, and it is characterized by the formation of all embryo cells from the apical cell (ca) of a two-celled proembryo. The only exception is that there is no formation of the radicle and/or cotyledons. The basal cell (cb) does not divide during the embryogenesis, gradually transforming into the uninuclear suspensor. Then the suspensor goes through three main stages: it starts with an unbranched cell within the embryo sac, followed by a branched stage growing into the integuments, and it ends with the cell death. The stage-specific development of the female gametophyte and embryo of D. nobile is discussed.

     

Embryology of the Theaceae - anther and ovule development of Camellia, Franklinia, and Schima

The anther and ovule development of Camellia, Franklinia, and Schima (Theaceae, Camellioideae) were observed. The three genera share the following embryological traits: anther wall formation of basic type, tapetum of glandular type, walls of endothecial cells with secondary thickening, and production of pseudopollen grains in connective, which are dispersed into pollen sacs at anthesis, ovule bitegmic-tenuinucellate, micropyle formed by inner integument alone, hypostase present, and both integuments generally five-to-seven cell layered. One autapomorphy of the Camellioideae found in the present study is the production of pseudopollen. The three genera surveyed differ with respect to the number of middle layers in the anther, the presence or absence of stomata on connective epidermis, morphology of pseudopollen, type of embryo sac formation, form of ovule, ovular vasculature, and the proliferation of ovular epidermis, etc. Among the three genera, Franklinia and Schima are presumed to be closer embryologically, and Schima possesses more numerous specialized features.     

ON THE ORIGINS OF THE OVULE AND CUPULE IN LYGINOPTERID PTERIDOSPERMS

Camp, Wendell H., and Mary M. Hubbard. (U. Connecticut, Storrs.) On the origins of the ovule and cupule in Lyginopterid pteridosperms. Amer. Jour. Bot. 50(3): 235–243. Illus. 1963.—The recently described Eurystoma angulare of the Lower Carboniferous with its naked, dichotomously branched, ovule-bearing branch truss may be taken conceptually as a starting point in a series of evolutionary reductions and modifications involving other known forms which ultimately led to the cupule surrounding the solitary ovule of later lyginopterids. It is postulated that the integuments of these ovules also were derived from dichotomously branched lateral trusses which immediately subtended the primitive megasporangia, but of less complexity than that which produced the cupule. Eurystoma indicates that ovules evolved independently of leaves; therefore, ovules cannot be thought of as having been derived from leaf tissues. Evidence is presented indicating that, although these pteridosperms produced ovules of considerable complexity, they did not bear seeds but dropped the pollinated ovules before fertilization. The already specialized organization of the ovules of the Lower Carboníferous pteridosperms indicates that the group must have originated in the Devonian. The structure of the Lyginopterid ovule is reinterpreted, indicating a basic similarity to that of the angiospermous ovule.     

Evolution and patterning of the ovule in seed plants

The ovule and its developmental successor, the seed, together represent a highly characteristic feature of seed plants that has strongly enhanced the reproductive and dispersal potential of this diverse group of taxa. Ovules encompass multiple tissues that perform various roles within a highly constrained space, requiring a complex cascade of genes that generate localized cell proliferation and programmed cell death during different developmental stages. Many heritable morphological differences among lineages reflect relative displacement of these tissues, but others, such as the second (outer) integuments of angiosperms and Gnetales, represent novel and apparently profound and independent innovations. Recent studies, mostly on model taxa, have considerably enhanced our understanding of gene expression in the ovule. However, understanding its evolutionary history requires a comparative and phylogenetic approach that is problematic when comparing extant angiosperms not only with phylogenetically distant extant gymnosperms but also with taxa known only from fossils. This paper reviews ovule characters across a phylogenetically broad range of seed plants in a dynamic developmental context. It discusses both well-established and recent theories of ovule and seed evolution and highlights potential gaps in comparative data that will usefully enhance our understanding of evolutionary transitions and developmental mechanisms.     

Ovule,fruit and seed development in Abolboda (Xyridaceae,Poales): implications for taxonomy and phylogeny

Xyridaceae belongs to the xyrid clade of Poales, but the phylogenetic position of the xyrid families is only weakly supported. Xyridaceae is divided into two subfamilies and five genera, the relationships of which remain unclear. The development of the ovule, fruit and seed of Abolboda spp. was studied to identify characteristics of taxonomic and phylogenetic value. All of the studied species share anatropous, tenuinucellate and bitegmic ovules with a micropyle formed by the inner and outer integuments, megagametophyte development of the Polygonum type, seeds with a tanniferous hypostase, a helobial and starchy endosperm and an undifferentiated embryo, seed coat derived from both integuments with a tanniferous tegmen and a micropylar operculum, and fruits with a parenchymatous endocarp and mesocarp and a sclerenchymatous exocarp. Most of the ovule and seed characteristics described for Abolboda are also present in Xyris and may represent a pattern for the family. Abolboda is distinguished by the ovule type, endosperm formation and the number of layers in the seed coat, in agreement with its classification in Abolbodoideae. The following characteristics link Xyridaceae to Eriocaulaceae and Mayacaceae, supporting the xyrid clade: tenuinucellate, bitegmic ovules; seeds with a tanniferous hypostase, a starchy endosperm and an undifferentiated embryo; and a seed coat with a tanniferous tegmen. A micropylar operculum in the seeds of Abolboda is described for the first time here and may represent a synapomorphy for the xyrids. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 144–154.