OVULE PACKAGING IN STOCHASTIC POLLINATION AND FERTILIZATION ENVIRONMENTS

The modular morphology of plants has important consequences for reproductive strategies. Ovules are packaged in discrete structures (flowers) that usually vary stochastically in pollen capture and ovule fertilization, because of the vagaries of pollen transfer by external agents. Different ovule packaging schemes may use limited reproductive resources more or less effectively, so that some number of ovules per flower may be optimal, given the prevailing probabilities of ovule fertilization. I derive a phenotypic model for ovule number per flower that maximizes the expected total ovule fertilizations on a plant when pollination and fertilization vary randomly among individual flowers. This model predicts that, except for small or inexpensive flowers, ovules should be “oversupplied” relative to the mean receipt of pollen tubes, so that pollen limitation of seed set should be common. Published data are congruent with this prediction. Additional hypotheses on the relation of ovule packaging to floral cost, plant size, and variance in pollen receipt are suggested by the model, but few data exist to evaluate these hypotheses.     

ADVENTIVE EMBRYOGENESIS IN CITRUS AND ITS RELATION TO POLLINATION AND FERTILIZATION

Cytological and histological studies of seeds from three facultative apomictic Citrus cultivars show that adventive embryos develop, as a rule, from the first few cell layers of the nucellus adjacent to the embryo sac in the micropylar half and occasionally from the chalazal end. The adventive embryos initiated in nucellar tissue away from the embryo sac and most of those initiated from the chalazal end of the nucellus do not develop beyond the one-celled stage. When two or more embryos are developing in the same seed, the successful development of a given embryo depends on its location in relation to access to nutrients from the endosperm. The presence of a zygote and triploid endosperm in seeds with adventive embryos, the abortion of seed when endosperm degenerates, and the lack of seed set without pollination indicate that pollination and fertilization are essential for in vivo adventive embryogenesis.     

OVULE ABORTION IN QUERCUS (FAGACEAE)

This study deals with the occurrence and relative abundance of four different types of abortive ovules in three species of Quercus. It was found that, contrary to previous literature, fertilization does not always occur in the abortive ovules. The most common type of abortive ovule is the one in which a normal embryo sac develops, yet fertilization does not occur. The absence of embryo sacs and the occurrence of empty embryo sacs account for abortion in other ovules. All types of abortive ovules can occur in the same ovary. It is proposed that all of the ovules that develop a normal embryo sac are potential seeds, but the first one to be fertilized suppresses the normal development of the others.     

穗花杉传粉和受精作用的研究

穗花杉于5月25日至6月15日(1980—1982)传粉,成熟花粉包括2个细胞。7月20日至29日受精。两者相隔约2个月。在花粉管和卵细胞中,分别具有许多特殊的核仁状颗粒,它是穗花杉属的明显特点之一。从传粉、受精和胚胎发育的特点来看,穗花杉和澳洲红豆杉属在红豆杉科中可能占居比较原始的位置,它们的演化顺序是:穗花杉属、澳洲红豆杉属、红豆杉属、白豆杉属和榧树属。而且,三尖杉科可能通过穗花杉属过渡到红豆杉科。     

POLLINATION AND SEED SET IN DIERVILLA LONICERA (CAPRIFOLIACEAE): TEMPORAL PATTERNS OF FLOWER AND OVULE DEPLOYMENT

Ovule development in Diervilla lonicera (Caprifoliaceae) is limited by insufficient pollination early in the blooming period and during extensive rainy periods. Production of flowers is skewed in time; an initial burst of flowering is followed by a long period of sparse flower production. Ovule number per flower increases through the blooming period. I discuss the interactions of fruit and seed set, ovule number, and bumble bee pollinator visitation patterns. When certain flowers have a higher a priori probability of successful pollination, it may be advantageous for plants to put more ovules in those flowers. Selective ovule deployment may be a general adaptive phenomenon that has received little attention.     

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.     

OVULE AND SEED DEVELOPMENT IN CERTAIN CACTI

Engleman , E. Mark . (U. of California, Davis.) Ovule and seed development in certain cacti. Amer. Jour. Bot. 47(6) : 460–467. Illus. 1960.—Ovules of Astrophytum myriostigma, Thelocactus bicolor, and Toumeya papyracantha are crassinucellate, possess a nucellar cap, and have a micropyle formed only by the inner of 2 integuments. Survival of the chalazal megaspore results in a monosporic gametophyte. A single layer of endosperm cells forms a cap over the radicle in a mature seed, as occurs in some of the Centrospermae. The characteristic hilum cup consists of the hilum and variable amounts of thickened cells of the funiculus which border a cuplike cavity. The hilum cup is associated with a funicular hump near the junction of the funiculus and the integuments. The funicular hump is similar in position to a swelling on the funiculus of some members of the Centrospermae.     

WIND POLLINATION,SELF-INCOMPATIBILITY,AND ALTITUDINAL SHIFTS IN POLLINATION SYSTEMS IN THE HIGH ANDEAN GENUS ESPELETIA (ASTERACEAE)

Altitudinal changes in breeding and pollination systems of tropical montane plants were studied in 13 species of Espeletia growing in the Venezuelan Andes from 2,000 to 4,300 m. Hand pollination tests showed that all species were strongly self-incompatible. The four species found only above 4,000 m had up to 10% median seed set in self-pollinated heads, which was significantly higher than the lower elevation species. Nine species were insect-pollinated, with large bees the major pollinator group. An endemic páramo hummingbird, Oxypogon guerinii, was an important visitor of E. schultzii in three populations examined. Experimental bagging experiments showed that the four high elevation species were wind-pollinated, further evidenced by the lack of pollinator visits and markedly different capitulum morphologies. Open-pollinated seed set in two wind-pollinated species, E. spicata and E. timotensis, was strongly dependent on the population's flowering density, which varied significantly from year to year. The shift from insect to wind pollination in Espeletia can be related to the low pollinator availability at high elevations in the Andes, protection of the capitula from snow and daily frosts, and the extremely long flowering periods of individual heads.     

VEGETATIVE DISPERSAL IN OXALIS CERNUA

Vegetative propagation and vegetative dispersal of Oxalis cernua were studied. O. cernua is a bulbous dicotyledonous plant. The shoot of a grown plant is divided into two distinct parts: a vertical stem growing upwards and bearing the feeding roots, and a rootless horizontal thin stem. This last is pulled by means of a large contractile root and carries along with it a series of small buds, which form new bulbs at the end of the season. Several new bulbs are formed along the vertical shoot as well. Thus, the propagation bulbs of O. cernua disperse along two axes, which are at right angles to each other.     

ADVENTIVE EMBRYOGENESIS IN CITRUS I. THE OCCURRENCE OF ADVENTIVE EMBRYOS WITHOUT POLLINATION OR FERTILIZATION

Anatomical studies of unfertilized undeveloped seeds from open- and control-pollinated fruits of ten facultative apomictic Citrus cultivars were carried out with the aid of light and epifluorescence microscopes. With or without pollination, adventive embryos autonomously developed at all positions in the nucellus in all cultivars. The adventive embryos initiated at the chalazal end of the nucellus were more vigorous than those initiated at the micropylar end. Because of the lack of endosperm and poor seed development, however, all adventive embryos within the unfertilized seeds terminated their development at the globular or early cotyledonary stages and were unable to germinate under natural conditions. The capability of unfertilized seeds to develop varied from species to species. Growth of the adventive embryos was dependent on nucellus size, but the growth rate of adventive embryos relative to nucellus size was different in different species. Neither pollination, fertilization nor subsequent zygote and endosperm development further stimulated adventive embryo initiation. Conversely, pollination and subsequent fertilization of other seeds in the same fruit slightly, but significantly, suppressed adventive embryo growth in the unfertilized seeds. These facts concerning adventive embryogenesis in unfertilized seeds indicate that neither pollination nor fertilization is essential for in vivo adventive embryogenesis and that normal endosperm is necessary for perfect development of adventive embryos initiated only in the micropylar half of the nucellus.     

POLLINATION AND POLLEN ENERGETICS IN MEDITERRANEAN ECOSYSTEMS

Most plants in Mediterranean ecosystems are insect pollinated, with pollen being the main reward to pollinators. The great majority of pollinators (70%) are bees and flies. We measured the energy content of pollen from 40 plant species in these ecosystems that represent abiotic and biotic pollination modes as well as the number of species of their pollinators. Pollen energy content correlates with pollinator diversity. Pollen of wind-pollinated plants contained less energy than that of insect-pollinated plants; there was no difference between insect-pollinated dicots and insect-pollinated monocots. The median date of flowering (from 1 January) estimated for each of the plant species did not vary significantly either with the number of pollinator species or with the energy content of pollen. The reasons for the differing values of pollen energy content are discussed; in particular, its relation to the type of pollen reserves, length of the flower style, and pollination enhancement.     

SPATIAL AND TEMPORAL VARIATION IN CHASMOGAMY AND CLEISTOGAMY IN OXALIS MONTANA (OXALIDACEAE)

Chasmogamous (CH) and cleistogamous (CL) flower production was investigated in natural populations of the perennial herb Oxalis montana in southern Quebec, Canada. Every 10–12 days during two flowering seasons, we recorded the reproductive output of approximately 2,000 plants distributed among five forest sites. The percentage of plants flowering, proportion of flowering plants producing CH and CL flowers, CH and CL flower number per flowering plant, and the proportion of flowers that are CL differed significantly among sites and corresponded to site differences in forest type and habitat quality. Unlike patterns in most CL species, however, reproduction by cleistogamy increased in sites with habitat conditions favorable to plant growth and sexual reproduction, and decreased in less productive sites. Chasmogamous and CL flower production increased with increasing plant size but plant size explained a significantly greater proportion of the variation in CL flower numbers. The percentage of flowering plants producing CH flowers decreased between years while the proportion of CL flowers increased at all sites during the second flowering season. The somewhat unusual aspects of reproduction in Oxalis may stem from its perennial habit which allows use of stored resources in adjusting the balance of cleistogamy versus chasmogamy in different environmental regimes.     

蝶兰胚珠的cDNA文库构建及其特异基因表达的研究

以蝶兰(Phalaenopsis“Mt.Kaala”cv SM9108)为材料,分别提取大孢子母细胞时期胚珠和成熟胚珠的PolyA RNA,反转录成cDNA,构建起两个cDNA文库。克隆筛选采用差异杂交法。从上述两个cDNA文库中,各选择一个筛选出的cDNA,对其在植物体不同器官和不同发育时期的胚珠内的表达进行了分析。结果表明该两个cDNA均为胚珠特异,并且分别在胚珠发育的特定时期表达。推测该两个cDNA的表达受胚珠内部的不同因子调控。     

DEVELOPMENT OF THE OVULE AND MEGAGAMETOPHYTE IN SAXIFRAGA HIERACIFOLIA

Wiggins , Ira L. (Stanford U., Stanford, Calif.) Development of the ovule and megagametophyte in Saxifraga hieracifolia. Amer. Jour. Bot. 46(10): 692–697. Illus. 1059.—Buds of Saxifraga hieracifolia collected in the vicinity of Point Barrow, Alaska, fixed, sectioned, and stained by standard methods, revealed that the archesporial cell in the ovule of this species is hypodermal and gives rise to the megaspore mother cell and a small number of parietal cells. Occasionally 2 megaspore mother cells occur within an ovule. Meiosis in the megaspore mother cell produces a linear tetrad of megaspores, the chalazal one of which normally gives rise to a monosporic, Polygonum-type megagametophyte. The polar nuclei fuse near the chalazal end of the megagametophyte and the antipodal cells disintegrate prior to fertilization. A distinct filiform apparatus and a marked lateral “spur” develop on each synergid. Vacuolation in the egg cell and in the synergids follows the usual pattern. Only a single integument surrounds the nucellus.     

CONE AND OVULE DEVELOPMENT IN SCIADOPITYS (TAXODIACEAE-CONIFERALES)

Ontogeny of seed cones of Sciadopitys, with special reference to the ovule-supporting structure, is studied in material collected in Japan and Massachusetts. Cones are initiated as lateral or terminal structures in summer and complete the formation of most organs before winter. Bract development is well advanced before ovule-supporting structures are initiated. Continued cone development involves the formation of a narrow ridge of tissue in the axil of each fertile bract. This ridge develops a series of nine (but up to 12) apical lobes in centrifugal order, each of which is the primordium of a future tooth on the ovuliferous scale. Ovules are initiated as outgrowths of the adaxial surface of each lobe so that there is a one-to-one ratio between lobes and ovules. Intercalary extension of the ovuliferous scale itself (distally) and the common base of the bract and ovuliferous scale (proximally) greatly extends the complex. The ovuliferous scale eventually exceeds the subtending bract and its apex becomes recurved. Bracts each have a single trace, but each ovuliferous scale has a pair of traces that proliferate distally to irrigate ovule and scale lobe. Intercalary growth results in recurvature of the ovule trace. The organization of the cone is directly comparable with certain Permian fossils. Sciadopitys also seems unique within the Taxodiaceae in its centrifugal development of the ovule-supporting complex.     

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.