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The paracarpous gynoecium in Capsella bursa-pastoris is characterized by postgenital fusion of two carpels into a single structure representing a false two-loculated ovary. The ovular primordium is initiated by periclinal cell divisions of both the subdermal and third layers of the placenta. The ovule is ana-amphitropous, medionucellate, funicular and bitegmic, with the micropyle formed by both integuments. During development the cells of the micropylar and middle nucellar zones degenerate and the persisting chalazal zone assumes a column shape (the postamento-podium). The integuments develop according to Dermal type and Variation C (author’s term). In the mature ovule the inner integument consists of three layers, and the outer integument is two-layered except on the abaxial side of the ovule where the integuments are more massive. At the two-nucleate megagametophyte stage the inner epidermis cells of the inner integument begin to divide periclinally. These divisions are followed by differentiation, giving rise to cells that differ in their form, structure, substance accumulation and participation in seed coat organization. The inner layer, consisting of cytoplasm-rich cells with marked radial expansion, represents the endothelium. The cells of the other layer become vacuolate and extend tangentially. They form the middle layer. The cells of the outer epidermis and middle layer are destroyed (in the latter only partly) during seed development. The endothelium becomes the endotegmen (pigment layer), composed of thick-walled cells that contain tannins and possibly lipids. The outer integument gives rise to the testa, composed of an epidermal mucilaginous layer and a sclerotic (mechanical) layer consisting of cells with thickened radial and inner tangential walls and containing starch. The hypostase is differentiated at the base of the nucellus and integuments in contact with the chalaza. The vascular bundle of the ovule reaches the hypostase, which is preserved also in the mature seed and represented by 3-5 layers forming a cup. The cells of the hypostase accumulate proteins and dextrins during the late stages of ovule development, and starch after fertilization. Later, at the early globular embryo stage, the cell walls begin to lignify, the cell contents showing tannin-like substances.
This study deals with the initiation and development of the integument in flowering plants. A classification including types and variations is proposed. The types (dermal, subdermal and dermal-subdermal) have different origins, that is, the integument may develop either from the dermal or subdermal or from both sets of initials. The characters of the variations comprise a complex of the following features: (1) the number of initials, (2) the sequence of their divisions and the mode of cell wall initiation at the first developmental stages, and (3) the participation of the initials in the formation of the various layers. The main evolutionary trends of the integument and ovule as a whole are considered.
The metabolites in the ovule come from the receptacle, ovary wall and placenta. They are accumulated in the chalaza and then translocated in reproductive and somatic structures via a system of specialized tissues. The hypostase is of great importance because it lies at the boundary of the chalaza with the nucellus and integuments, and contacts the vascular bundle. In the megagametophyte, the cell walls, which have many outgrowths, receive metabolites from the hypostase via podium and postament tissues. Histochemical data on accumulation of proteins, polysaccharides, lignin and tannins in tissues during ovule and seed development are presented. The dynamics of these substances and the character of metabolism in tissues are suggested as indicators of metabolite flow.
A study of Gentiana cruciata L. (Gentianaceae), Gymnadenia conopsea (L.) R.Br. (Orchidaceae) and Luzula pedemontana Boiss. et Reut. (Juncaceae) showed differences in the number and characteristics of critical stages in ovule and seed development. The shared critical stages explain the general direction of the formation of reproductive structures and surrounding tissues. The taxon-specific critical stages may have different implications in a given species: they may (1) verify that the ovule belongs to a specific type, (2) indicate their lability in different taxa with the same ovule type, or (3) coincide in species with various ovule types.
Ovule and seed morphogenesis in Vaccinium myrtillus was studied here for the first time. The gynaeceum is paracarpous. The ovules are developed at sutural-angular placentae. Periclinal divisions of the subepidermal cells of the placenta usually precede formation of the ovular primordia. The ovules are ana-campylotropous, medionucellate, unitegmic, leptochalazal and sessile. The initials and dynamics of different ovule structures were determined. The chalazal zone of the nucellus transforms into the postamento-podium. The integument is of dermal origin, with a typical endothelium. The ovule has a hypostase, which is initiated with the beginning of ovular primordium differentiation. It is cup-shaped in the mature ovule and consists of 2-3 layers. Their cell walls are not lignified. The nucellus, integumental parenchyma, endothelium and hypostase do not persist in the mature seed. The latter is exotestal. The embryo sac develops according to the Polygonum type. The endosperm is cellular, with micropylar and chalazal haustoria. The mature embryo is large and well differentiated into organs.
Comparison of maize mutants pam1, mac1 and normal plants (variety Belaya noch’) showed that the formation of female reproductive organs is very similar between them. One afunicular ovule develops in an ovary; it is ortho-campylotropous and bitegmic, with a hypostase. The postament, podium and nucellar cap are differentiated in the nucellus. The integuments, epidermal in origin, arise from a common initial zone, but their differentiation is separated in time. In the subepidermal layer of the apical part of the primordium, the complex of initial cells is distinguished. Some of them differentiate as archesporial cells. In normal plants and pam1, only one of them transforms into the megasporocyte, while up to 6-8 do in mac1. Polygonum-type embryo sacs are formed. In pam1 the majority of megasporocytes do not undergo meiosis, while the surrounding ovular tissues continue their development. Analysis of the distribution of polysaccharides and proteins revealed similarities in their spatial and temporal coordination. The differences are quantitative, and probably conditioned by the number of megaspores and embryo sacs.
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