Persistent Synergid Research Articles

Fertilization of Crocus sativus L. ovules and development of seeds after stigmatic pollination with C. thomasii Ten. pollen

Abstract Fertilization and seed development were studied in Crocus sativus L., after pollination with Crocus thomasii Ten. Fertilization was porogamous and resulted in the formation of zygote and primary endosperm nucleus; it was accompanied by degeneration of the receptive synergid and followed by degeneration of the antipodals. The endosperm developed precociously by free nuclear division, occuping a parietal position; subsequently it filled the central vacuole and became cellularized. The zygote divided after a quiescence period by formation of a transversal wall; its division was accompanied by degeneration of the persistent synergid. The embryo developed from the apical cell and, at maturity, was a cylindrical structure showing histological differentiation of the shoot apex. The seed coat was formed from both integuments. Both embryogeny and development of the endosperm and seed coat were entirely similar to those of Crocus thomasii.

Seed development of Ornithogalum dubium, with special reference to fertilization and the egg apparatus

The ovule of Ornithogalum dubium is anatropic, bitegmic and crassinucellate. The integuments are biseriate. Starch grains occur in the outer integument, the funicule and in the intercellular space surrounding the embryo-sac. The embryo-sac development is of the Polygonum type. The synergids are conspicuous and contain a large filiform apparatus. Fertilization is porogamic and occurs 24 h after pollination. The pollen tube enters the egg apparatus in the region of an intercellular space between the micropylar ends of the synergids. Sperm cells in the pollen tube appear to be released through small protuberances of the pollen tube. Migration of the sperm cell to the egg cell is possibly through a gap in the wall between the egg cell and the receptive synergid. Changes in the synergids occur only after a pollen tube has entered the receptive synergid. Changes occurring in the persistent synergid suggest that this cell may be involved in the nutrition of the pro-embryo up to the eight-cell pro-embryo stage. Syngamy is of the post-mitotic type. The first division of the zygote is transverse. The basal cell gives rise to the multicellular suspensor, and the terminal cell to the embryo. The endosperm is of the Helobial type.

Some ultrastructural aspects of the pollen transmission of barley stripe mosaic virus in barley

To study the pollen transmission of barley stripe mosaic virus (BSMV) in barley with the transmission electron microscope, ultrathin sections of ovaries of pollinated pistils were examined. Healthy pistils were cross-pollinated with virus-infected pollen and collected at various intervals of time after pollination. When thin sections of the ovaries were viewed, virus particles were detected in specific sporophytic and gametophytic cells during critical developmental stages of pollination, fertilization, and embryogenesis. Before fertilization, BSMV particles were seen in the pollen tube between the integument and ovary wall, and in pollen tube discharge within the degenerating synergid. During and after fertilization, virus particles were found not only in the pollen tube and its discharge but also in the zygote, endosperm, persistent synergid, and nucellus. During embryogenesis, BSMV particles were very evident in the embryo, integument, and ovary wall. Some particles were scattered throughout the cell cytoplasm, while others were in mono- or multi-layer aggregates within the cytoplasm. Many particles were associated with spindle or cytoplasmic microtubules or with abnormal plastids. The prevalence of virus particles in the cells associated with sexual reproduction suggests that the nucleoprotein particle form of the virus plays some role in the pollen transmission of BSMV in barley.

Early Development of the Macadamia Ovary

Macadamia integrifolia ovaries were sampled for light microscopy from anthesis to 8 weeks after flowering. Both ovules increased in size following anthesis but only one was normally fertilized. Fertilization of one ovule appeared to inhibit fertilization of the other. The unfertilized ovule was eventually crushed by the fertilized ovule which developed a globular embryo with a short suspensor. Division of the endosperm preceded division of the embryo, and cell formation in the initially free nuclear endosperm progressed by the development of cell walls inwards from the embryo sac wall at the micropylar end of the embryo sac only. Thickenings developed on the walls of the embryo sac, persistent synergid and embryo and a hypostase developed at the base of the nucellus between the cavity occupied by the embryo sac and the integument. Both specializations are suggested to be important in the nutrition of the developing embryo. Ovaries which abscissed 4-5 weeks after anthesis had a similar anatomy to ovaries retained on the tree. Some embryos showed signs of degeneration but this was not considered to be the cause of abscission.

POLLEN TUBE‐SYNERGID INTERACTIONS IN PROBOSCIDEA LOUISIANICA (MARTINEACEAE)

The ultrastructure of the synergids of Proboscidea louisianica was investigated from just before fertilization until 48 hr after pollination. It was found that the cytoplasm of one synergid consistently begins to degenerate before arrival of the pollen tube at the embryo sac, and that it is always this synergid which receives the pollen tube tip and its discharge. The other synergid (persistent synergid) remained unchanged throughout the study period. Polysaccharide vesicles of pollen tube origin were observed fusing with the pollen tube wall as well as contributing to cell wall formation of the degenerate synergid. In one ovule (48 hr after pollination) two pollen tubes had entered and grown the length of the micropyle, but only the first tube penetrated the degenerate synergid and discharged normally. The second pollen tube was abutting against the persistent synergid, but had not entered or discharged. In another exceptional case (18 hr after pollination), a pollen tube had grown the length of the micropyle, but did not discharge, or enter either synergid. Both synergids of this ovule were observed to be completely intact. It is concluded that synergid and pollen tube cytoplasmic degeneration is the result of a very specific interaction between these two cells and that this degeneration is probably a prerequisite for normal pollen tube entrance and discharge into the embryo sac, and for male gamete transfer to the egg and central cell.

A study of the mature megagametophyte of Stipa elmeri

In Stipa elmeri Piper & Brodie ex Scribn., the pollen tube enters at the filiform apparatus of the degenerated synergid. The degenerated synergid has electron-dense cytoplasm in which organelles are not discernible. All other cells of the mature megagametophyte have nuclei, endoplasmic reticulum, plastids, mitochondria, dictyosomes, and vacuoles. Starch is found in the persistent synergid (in minute quantities), egg, and central cell. Lipids occur in the persistent synergid, central cell, and antipodals. The filiform apparatuses of the two synergids are hypothesized to perform different functions. In the degenerated synergid, the filiform apparatus serves to increase the surface area of the plasma membrane and thereby to offer a large area for pollen-tube-growth-directing compounds to diffuse out of the synergid. In the persistent synergid, the filiform apparatus is part of a suite of features which indicate that the persistent synergid is involved in the transference of materials into the megagametophyte. Another possible function of the persistent synergid is to aid in establishing the polarity of the egg. The pollen grain and tube have distinctive polysaccharide spheres that serve to delimit the pollen tube cytoplasm after discharge into the degenerated synergid. Associated with the degenerated synergid are bodies of dense materials as seen under electron microscopy, and bodies of RNA and protein as determined histochemically. These are probably the same thing and come from the degenerating synergid. The antipodals are the most cytologically active cells of the megagametophyte. They have some features which are characteristic of transfer cells and possibly function in the transference of materials into the megagametophyte. Other studies (Brink and Cooper 1944) have indicated that grass antipodals are involved in the control of endosperm development. The active cytoplasm of the antipodals may reflect the synthesis or transference of growth-controlling substances.

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

The degeneration of one synergid denotes the initiation of embryo and endosperm development in the embryo sac of sunflower Helianthus annuus L. The other synergid, the persistent synergid, is present until the late globular stage of embryogenesis. The primary endosperm nucleus divides before the zygote nucleus forming a coenocytic nuclear endosperm. When about eight endosperm nuclei are present during the early globular stage of embryogenesis, endosperm wall formation starts at the micropylar end of the embryo sac. The walls continue to grow toward the chalazal end of the embryo sac apparently as a result of the activity of Golgi located at the tips of the growing walls. Most endosperm wall formation is not associated with a mitotic spindle apparatus in sunflower. The suspensor of the embryo consists of a large basal cell during the proembryo stages, a single row of cells during the early globular stages, and at the late globular stage a double tier of cells near the radicle end of the embryo and a single row at the micropylar end of the embryo sac. Occasionally embryo development occurs in the absence of endosperm when only single fertilization has taken place. The development and nutritional implications of post-fertilization events in the sunflower embryo sac are discussed.

FINE STRUCTURE AND COMPOSITION OF THE EGG APPARATUS BEFORE AND AFTER FERTILIZATION IN QUERCUS GAMBELII: THE FUNCTIONAL OVULE

A study of the egg apparatus of Quercus gambelii was made at both the light and the electron microscope levels. This investigation was concerned primarily with the changes that occur in these cells before and after the process of fertilization and what role, if any, is played by the synergids in this phenomenon. The synergids before fertilization are, on the basis of ultrastructure, healthy, intact, functional cells. They have numerous mitochondria, dictyosomes, endoplasmic reticulum, ribosomes, and a typical nucleus. A prominent filiform apparatus is present, but the cell wall only extends a short distance around the micropylar end of the cells. Just before fertilization, one of the synergids degenerates. This is the synergid that receives the pollen tube and its discharge, including both male gametes. Dictyosomes increase in number and activity in the other synergid (persistent synergid) after fertilization. Eventually a complete cell wall forms around both of the synergids. No plasmodesmata are present in these walls. The egg has numerous mitochondria, dictyosomes, endoplasmic reticulum, and ribosomes, both free in the cytoplasm and attached to the endoplasmic reticulum. Lipid bodies are characteristic of this cell. A cell wall is present only around the micropylar end of the egg. After fertilization, little change occurs in the zygote. The number and activity of the dictyosomes increase, apparently in correlation with cell wall formation. The number of lipid bodies increases. The zygote is approximately the same size as the egg. Plastids are scarce, and starch grains are typically absent from all cells of the egg apparatus. It is suggested that the synergids function in the secretion of chemotropic substances that guide the growth of the pollen tube. Comparisons are made between the egg apparatus of Quercus gambelii and that of the other plants studied thus far.

Fine Structure and Composition of the Egg Apparatus Before and After Fertilization in Quercus gambelii: The Functional Ovule

A study of the egg apparatus of Quercus gambelii was made at both the light and the electron microscope levels. This investigation was concerned primarily with the changes that occur in these cells before and after the process of fertilization and what role, if any, is played by the synergids in this phenomenon. The synergids before fertilization are, on the basis of ultrastructure, healthy, intact, functional cells. They have numerous mitochondria, dictyosomes, endoplasmic reticulum, ribosomes, and a typical nucleus. A prominent filiform apparatus is present, but the cell wall only extends a short distance around the micropylar end of the cells. Just before fertilization, one of the synergids degenerates. This is the synergid that receives the pollen tube and its discharge, including both male gametes. Dictyosomes increase in number and activity in the other synergid (persistent synergid) after fertilization. Eventually a complete cell wall forms around both of the synergids. No plasmodesmata are present in these walls. The egg has numerous mitochondria, dictyosomes, endoplasmic reticulum, and ribosomes, both free in the cytoplasm and attached to the endoplasmic reticulum. Lipid bodies are characteristic of this cell. A cell wall is present only around the micropylar end of the egg. After fertilization, little change occurs in the zygote. The number and activity of the dictyosomes increase, apparently in correlation with cell wall formation. The number of lipid bodies increases. The zygote is approximately the same size as the egg. Plastids are scarce, and starch grains are typically absent from all cells of the egg apparatus. It is suggested that the synergids function in the secretion of chemotropic substances that guide the growth of the pollen tube. Comparisons are made between the egg apparatus of Quercus gambelii and that of the other plants studied thus far.

Embryo sac and early ovule development in Oryzopsis miliacea and Stipa tortilis

The ovules of Stipa tortilis and Oryzopsis miliacea are hemianatropous, bitegmetic, and pseudocrassinucellate (sensu Davis 1966). The hemianatropous shape of the ovule is the result of characteristic patterns of cell division and enlargement in the chalazal area and areas alongside the embryo sac. Embryo sac development in both is Polygonum-type and both have proliferating antipodals. Endosperm is nuclear, although in O. miliacea it is atypical in that nuclear division is synchronous within one portion of the embryo sac, e.g. micropylar, but not synchronous between different portions of the embryo sac, e.g., micropylar and chalazal. Differences in ovule initiation, persistence of the outer integument, fate of the inner integument, nature of the nucellus, shape of the embryo sac, nature of the synergids, cytoplasm of the egg, polar nuclei, and endosperm exist between these two taxa. Both synergids of O. miliacea undergo changes before fertilization and one degenerates before fertilization. The pollen tube enters the embryo sac at the base of the persistent synergid. There is presently insufficient embryological data to permit meaningful speculation on relationships between Stipa and Oryzopsis. Embryologically, Stipa and Oryzopsis are festucoid grasses, as much other evidence indicates. Embryo sac development in the Gramineae is more similar to that of the Restionaceae than to that of the Cyperaceae. This is in contradiction to recent speculations on the relationships of the Gramineae.