Development of Male and Female Gametophyte in Symphytum orientale L. (Boraginaceae)

The presented study elucidates developmental features of male and female gametophyte in Symphytum orientale L. (Boraginaceae). Flower buds were collected from natural habitats (İstanbul-Turkey) in March-April. The anthers of S. orientale were tetrasporangiate with a persistent epidermis and its wall development conformed to the Dicotyledonous type. The endothecium developed fibrous wall thickenings. The middle layer was short-lived and the cells of the glandular tapetum became double nucleated. In the microspore mother cells cytokinesis originated simultaneously after meiosis and the microspore tetrads were tetrahedral or isobilateral. The mature pollen grains were 3-celled when spread to environment. The ovule was hemianatropous, unitegmic and tenuinucellar. The megaspore mother cell functioned directly from the archesporial cell. Polygonum type embryo sac developed from the chalazal megaspore of a linear tetrad generated after meiosis. The synergids were pyriform and the polar nuclei fused shortly before fertilization. The antipodal cells were large and persistent at the stage of globular embryo. Embryogenesis followed the Chenopodiad type and the endosperm represented cellular type. The detailed embryological data of Symphytum orientale L. (Boraginaceae), which is a herbaceous and perennial flowering plant will improve our knowledge of its reproductive behaviour, and provide to comprehend taxonomic connection with related taxa within the Symphytum/Boraginaceae.

Female gametophyte and pollen tube of Epilobium palustre L.

The monosporic, tetranucleate embryo sac of <i>Epilobium palustre (Onagraceae)</i> develops from the micropylar megaspore in a linear tetrad. In mononucleate embryo sacs a peculiar chromatic structure associated with a nucleolus appears in the nucleus. This structure seems to be formed by fibrillar material and is not visible in the subsequant stages of development. A large amount of rough ER cisternae occurs in the late mononucleate stage, during the binucleate stage their contents become optically dense. It the early tetranucleate stage the amount of ER is small, it increases again in the developing synergids and central cell. Numerous amyloplasts present in the mononucleate embryo sac loose their starch grains and some are transformed into cup-shaped plastids or proplastids. They are passed on to each of the embryo sac cells. The growth of the pollen tube ceases immediately after the penetration through the filiform apparatus of a synergid. At the apex of the tube a pore is formed. At the last stages of the penetration the apical part of the pollen tube becomes separated by a transverse partition from the distal part of the tube. The contents of the both parts differ in their internal structure. The distal part contains cytoplasm with numerous organoids, while the apical part is mainly filled with spherical bodies.

Female gametophyte development and embryogenesis in Taxus baccata L.

Crassinucellate ovules are initiated in <em>Taxus</em>, directly from the shoot apex. The rudimentary pollen chamber is formed in the nucellus. A linear tetrad of megaspores with a functional chalazal megaspore is formed. A free-nuclear stage is charac-teristic at the beginning of megagametophyte development. Archegonia without ventral canal cell are solitary or in complexes. The embryo has a very long suspensor even after maturation. Two types of polyembryony have been revealed: i) embryogenic redifferentiation of suspensor cells and ii) cleavage of embryonic region in the early embryo. In the northern temperate climate of St. Petersburg one month delay in development of reproductive structures has been noted.

Mating Within the Meiotic Tetrad and the Maintenance of Genomic Heterozygosity

Mating among the products of a single meiosis (automixis or meiotic parthenogenesis) is found in diverse groups of plant, animal, and fungal taxa. Restoration of the diploid stage is often strictly controlled and brings together products separated at the first meiotic division. Despite apparent similarities to diploid selfing, the theoretical prediction is that heterozygosity should be maintained on all chromosomes when it is linked to the centromeres and thus also segregates at the first meiotic division. Using the fungus Microbotryum, we directly test this prediction by linear tetrad analysis. The patterns of meiotic segregation for chromosome size variation (electrophoretic karyotypes) and PCR products (AFLP procedures) were determined for Microbotryum lineages native to North America and Europe. Our data reveal a surprisingly dynamic genome that is rich in heterozygosity and where size-dimorphic autosomes are common. The genetic variation agrees with the prediction of centromere-linked heterozygosity. This was observed to the greatest extent in the lineage of Microbotryum native to North America where there was consistent first-division segregation and independent assortment of multiple linkage groups. The data also show properties that distinguish the fungal sex chromosomes from the autosomes in both lineages of Microbotryum. We describe a scenario where the mating system of automixis with first-division restitution is the result of feedback mechanisms to control exposure of genetic load.

The embyology and relationship of the Byblidaceae

The relationships of the Byblidaceae have been the subject of a number of recent molecular phylogenetic studies where their traditional relationships with the Roridulaceae and other members of the Rosidae have been overturned in favour of affinities with the Asteridae, in particular to the Lentibulariaceae in the Lamiales. Although the embryological relationships between these families were the subject of an earlier study, the data for the Byblidaceae were incomplete. The family has tetrasporangiate, bilocular anthers with a glandular bi-nucleate tapetum. Formation of the anther wall appears to be of the Dicotyledonous type, and the anthers have ephemeral middle layers and apical fibrous thickenings. Simultaneous microsporogenesis results in tetrahedral tetrads of bi-nucleate pollen grains. The ovules are anatropous, unitegmic and tenuinucellate. Megasporogenesis is direct from the archesporal cell, which in Byblis liniflora Salisb. divides to produce a linear tetrad from which a chalazal megaspore is derived (contrary to a report of a micropylar megaspore in B. gigantea Lindl.). Megagametogenesis conforms to the Polygonum type, and the antipodal cells are persistent. The endosperm is ab initio Cellular and there are well developed micropylar and chalazal haustoria. Embryogenesis appears to conform to the Onagrad type noted for several of the other members of the Lamiales s.l., including the Lentibulariaceae, to which the Byblidaceae have been related in recent molecular studies.

Anther and Ovule Development in Tasmannia (Winteraceae)

Three species of Tasmannia R.Br. ex DC., T. glaucifolia, T. insipida and T. stipitata are studied. The anther is tetrasporangiate and its waU development conforms to the Basic type. The tapetum follows the secretory type of development. Cytokinesis in the microspore mother cells is simultaneous but an evanescent cell plate is present at telophase I and anaphase I1 during meiosis. Pollen tetrads are permanent and tetrahedral. The mature pollen is anaulcerate, reticulate and 2-celled. The ovule. is anatropous, bitegmic and crassinucellate. The micropyle in T. stipitata and T. Glaucifolia is formed by the inner integument only whereas in T. insipida it is formed by both the integuments and is zigzag in outline. Meiosis in the single megaspore mother cell produces a linear or T-shaped megaspore tetrad in T. stipitata and T. glaucifolia but only a linear tetrad in T. insipida. Female gametophyte development is of the monosporic Polygonum type. Fertilisation is porogamous; triple fusion and syngamy occur simultaneously.

Megasporogenesis in normal and a synaptic-mutant (sy-2) of Solanum commersonii Dun.

Nomarski differential interference contrast microscopy was used to study megasporogenesis in intact ovules of Solanum commersonii Dun., following staining with Mayer's hemalum and clearing with methyl salicylate or cedarwood oil. Previous studies in Solanum have observed the Polygonum type of megasporogenesis, in which a linear tetrad of megaspores is formed. The three micropylar megaspores then degenerate, leaving the chalazal megaspore to divide mitotically to form the egg sac. Contrary to expectation, only 30% of the observed sporads within the same ovary were tetrads. Triads, including some with one deteriorating cell at the micropylar end, were the predominant form. Many dyads, which are an intermediate stage in megasporogenesis, were found with one cell prematurely deteriorating. These observations can be explained if the micropylar daughter cell of the dyad stage, which formed after telophase I, began deteriorating before the second meiotic division, such that it never underwent the second division. The chalazal daughter cell would still undergo the second meiotic division, followed by death of the new micropylar megaspore.Key words: Solanum, potato, megasporogenesis, S. commersonii, ovule, clearing.

Ordered Linear Tetrads Are Produced by the Sporulation of Newly Formed Zygotes of Saccharomyces cerevisiae

ABSTRACT Diploid Saccharomyces cerevisiae strains normally sporulate to produce tetrahedral unordered asci containing four spores (tetrads). We report that when newly formed zygotes are subjected to the same sporulation conditions, they form predominantly linear ordered tetrads. We show that the two spores from each end of such a linear tetrad invariably contain nonsister centromeres. Spore viability, recombination and independence of centromere segregation appear unaffected.

Sexual reproduction of mountain hemlock (Tsuga mertensiana)

Meiosis of pollen mother cells begins in October of the year in which cones are initiated. They reach pachytene then become dormant until the next March. Meiosis is complete and the winged pollen mature by mid-June. Meiosis of the megaspore mother cell occurs in May, forming a linear tetrad of megaspores. The female gametophyte undergoes free nuclear division at pollination in mid-June. No pollination drop is present; rather, the pollen adheres to the sticky, splayed edge of the micropyle, where it germinates and pollen tubes grow toward the nucellus. The nucellus elongates into the micropylar canal, forming a nucellar beak, which makes contact with the pollen tubes. Several pollen tubes penetrate the nucellus.At the time of fertilization early in August, each ovule contains two to four aichegonia each having two to four neck cells in one tier. Pollen tubes penetrate the neck cells and two male gametes are formed. The ventral canal cell breaks down and fusion occurs in the center of the archegonium. Four free nuclei form and migrate to the base of the archegonium. cell walls form, and a 16-celled proembryo develops. Both simple and cleavage polyembryony occur. Rosette cells divide but do not form complex embryos. The embryo and seed are mature in October and the cones dry and open during October and November. Mature cones averaged 70 seeds, of which 46% were filled.Reproduction in mountain hemlock (Tsuga mertensiana (Bong.) Carr.) is similar to that in other species of Tsuga except for the presence of winged pollen. Any attempt to place the species in the genus Picea or place it as a hybrid midway between Picea and Tsuga is unfounded based on all of the more-conservative reproductive and embryological characteristics.

The development of the embryo sac of sunflower Helianthus annuus before fertilization

The megaspore mother cell of sunflower Helianthus annuus L. undergoes two meiotic divisions to form a linear tetrad of haploid megaspores. The chalazal megaspore increases in size while the other megaspores and the nucellus degenerate such that the integumentary tapetum is adjacent to the embryo sac. Mitotic divisions occur forming the coenocytic two- and four-nucleate embryo sacs and the seven- or eight-nucleate six-celled embryo sac. Electron-microscopic observations suggest that the antipodals are very active synthetically but start degenerating before fertilization. Similarly the synergids are also apparently very active synthetically before fertilization as judged by the presence of extensive regions of dilated rough endoplasmic reticulum and many Golgi bodies and associated vesicles. The egg cell is characterized by the presence of many free ribosomes and small undifferentiated plastids. The central cell contains many circular strands of rough endoplasmic reticulum, lipid droplets, and large clusters of apparently active Golgi; it is a transfer cell resulting from the presence of embryo sac wall ingrowths. The development and the possible nutritional interrelationships of the megagametophyte and surrounding tissues are discussed.