Order of fertilization within the ovary in Phaseolus coccineus L. (Leguminosae)

Summary Phaseolus coccineus typically has six linearly arranged ovules per ovary. The three ovules near the stylar end of the fruit (positions one, two, and three) are more likely to produce mature seeds, to produce heavier seeds, and to produce more vigorous progeny than the ovules in positions near the peduncular/basal end of the fruit (ovule positions four, five, and six). We conducted a series of field experiments designed to supplement our understanding of the mechanisms determining these position effects. We found that approximately 98% of the ovules in 752 fruits were fertilized — about 0.6% of the stylar ovules were not fertilized, whereas 3.2% of the basal ovules were unfertilized. Moreover, we found that only about 49% of the ovules in these 752 fruits produced mature seeds. Over 60% of the stylar ovules produced mature seeds, whereas only 37% of the basal ovules produced mature seeds. Consequently, the proportion of fertilized ovules cannot explain the differences in seed maturation among the ovule positions. We found that after 6.5 h most of the fertilized ovules were located in the stylar ovule positions, and that there were no fertilized ovules in ovule positions five and six, indicating that the stylar ovules are fertilized first. When only the fastest growing pollen tubes were permitted to enter the ovary (due to exision of the style), only the ovules at the stylar end were fertilized, indicating that the ovule positions that are fertilized first are indeed fertilized by the fastest growing pollen tubes.On leave from the Escuela de Biologia, Universidad de Costa Rica, Cuidad Universitaria Rodrigo Facio, San Jose, Costa Rica, Central America     

Unfertilized ovules of Epilobium obcordatum (Onagraceae) continue to grow in developing fruits

To determine whether unfertilized ovules continue to grow when in an ovary containing fertilized ovules, we measured ovule lengths in developing fruits of Epilobium obcordatum that were harvested 4, 5, 8, and 10 d post pollination. We found that unfertilized ovules that were in the presence of fertilized ovules continued to grow and that there was a broad range of overlap in their sizes at all sampling times. This effect was found for two types of unfertilized ovules that occur throughout the length of the ovary: normal, unfertilized ovules, apparently bypassed by pollen tubes; and sterile ovules lacking an embryo sac. In addition, there is a position effect within developing fruits. Both fertilized and unfertilized ovules are larger at the stylar end. In six samples resulting from pollination with a single pollen tetrad, a total of 18 embryos were found, and the effect on unfertilized ovules, greatest at the stylar end, diminished with distance from the ovules with embryos. Our results are consistent with the interpretation that diffusible hormones produced by developing seeds cause nearby unfertilized ovules to grow. We conclude that caution is necessary when attempting to infer ovule fertilization histories from the appearances of ovules in developing and mature fruits. What are often inferred to be aborted seeds, in many cases, may not be seeds at all. They may be enlarged, unfertilized ovules.     

Multinucleate storage cells in Douglas-fir (Pseudotsuga menziesii (Mirbel) Franco) and the effect of seed parasitism by the chalcid Megastigmus spermotrophus Wachtl

Megagametophytes of Douglas-fir (Pseudotsuga menziesii (Mirbel) Franco) accumulated storage products following fertilization. As megagametophytes matured, the number of nuclei per cell rose, resulting in syncytial storage cells. Studies carried out on trees in France and Canada confirmed that such previously unreported, free nuclear cells were a normal part of late megagametophyte development. Unfertilized megagametophytes showed that some binucleate cells before degeneration resulted in empty seed. Insect parasitism prevented megagametophyte abortion in unfertilized ovules. Oviposition by a torymid chalcid wasp (Megastigmus spermotrophus Wachtl) early in megagametophyte development resulted in normal megagametophyte development. Around the time of plant egg maturation, binucleate and trinucleate cells were observed. As megagametophytes matured, multinucleate mature storage cells rich in proteins, lipids and starch were formed. The insect was able to induce identical nuclear behaviour in infested, unfertilized megagametophytes, as that of uninfested, fertilized megagametophytes.     

POLLINATION OF THE ARROYO WILLOW,SALIX LASIOLEPIS: ROLE OF INSECTS AND WIND

We studied the relative roles of insects and wind as pollen vectors for the arroyo willow, Salix lasiolepis Benth. through insect exclosure studies at two elevational sites. Insect pollination resulted in production of 99.9% of seeds at a low elevation site (2,130 m) and at a high elevation site (2,450 m) with wind playing an exceedingly small role. Few pollen grains reached individual pistillate catkins despite the presence of abundant airborne willow pollen near male plants. In flowers that showed evidence of receipt of wind transferred pollen, due to their swollen appearance, we found only 1.74 fertilized ovules per flower and that 90.9% of these fertilized ovules were aborted by the maternal plant. In contrast, flowers open to insect visitation possessed 7.00 fertilized ovules per flower, 83.3% of which became filled seeds. We tested for the existence of apomictic seed production through use of closed bags that excluded both insect-and wind-transferred pollen but found no evidence of seed produced in this treatment. Additional control studies documented that wind treatment bags did not impede pollen flow while neither wind or closed bags influenced seed set in our treatments.     

Effect of ovule position within the pod on the probability of seed production in Bauhinia ungulata (Fabaceae)

BACKGROUND AND AIMS: It has been claimed that ovules linearly ordered within a fruit differ in their probabilities of reaching maturity. This was investigated by studying the effect the position of an ovule within the pod has on seed abortion and seed production in Bauhinia ungulata. METHODS: Fruits collected during the dry seasons of 1999, 2000 and 2001 were opened, and the number, position and status of each ovule within the fruit were recorded. A GLM model was used to assess the effects of population, tree identity and ovule position within the pod on ovule fertilization, seed abortion, seed damage and seed maturation in two populations of B. ungulata. KEY RESULTS: Nearly 30% of the ovules were not fertilized in 1999; this percentage dropped to 5% the following two years. Seed abortion (50%) and seed damage (15%) were the same every year during the study period. Only 15% of the initial ovules developed into mature seeds in 1999; this value increased to 35% in 2000 and 2001. However, seed survivorship was dependent on the position of the ovule within the pod; non-fertilized and early aborted ovules were found more often near the basal end of the ovary. The frequency of seed damage was not affected by position. Mature seeds were found mainly in the stylar half of fruits, where ovules are likely to be fertilized by fast pollen tubes. CONCLUSIONS: The pattern of seed production in B. ungulata is non-random but is dependent upon the position of the ovule within the pod. The results suggest that the seeds produced within a fruit might differ in their vigour.     

Regeneration of Fertile Barley Plants from Mechanically Isolated Protoplasts of the Fertilized Egg Cell

A simple procedure is described for the mechanical isolation of protoplasts of unfertilized and fertilized barley egg cells from dissected ovules. Viable protoplasts were isolated from ~75% of the dissected ovules. Unfertilized protoplasts did not divide, whereas almost all fertilized protoplasts developed into microcalli. These degenerated when grown in medium only. When cocultivated with barley microspores undergoing microspore embryogenesis, the protoplasts of the fertilized egg cells developed into embryo-like structures that gave rise to fully fertile plants. On average, 75% of cocultivated protoplasts of fertilized egg cells developed into embryo-like structures. Fully fertile plants were regenerated from ~50% of the embryo-like structures. The isolation-regeneration techniques may be largely genotype independent, because similar frequencies were obtained in two different barley varieties with very different performance in anther and microspore culture. Protoplasts of unfertilized and fertilized eggs of wheat were isolated by the same procedure, and a fully fertile wheat plant was regenerated by cocultivation with barley microspores.     

Structural aspects of ovule and seed development and nonrandom abortion inMelilotus officinalis (Fabaceae)

Summary Only one ovule matures into a seed inMelilotus officinalis. Although eight ovules form within an ovary, only the basal ovule develops into a mature seed, whereas the other ovules degenerate. The investigation of ovule and seed structure at different developmental stages and a comparison of quantitative characters of differently fated ovules within an ovary were undertaken by light, phase contrast, and fluorescence microscopy. In this species, campylotropous ovules develop simultaneously on marginal placentae in an apocarpous unilocular gynoecium. Megasporo- and megagametogenesis proceed normally and are completed in bud. The maturation of the Polygonum type embryo sac takes place after the flower opens. Shortly before fertilization, synergids show signs of degeneration in all ovules. At this stage, neither the structure nor the sizes of ovules within one ovary differ significantly. In spite of this, only the basal ovule develops into a seed. Rarely, one of the upper-situated ovules or the basal and another ovule mature into seeds. Seed enlargement is insignificant until the stage when globular embryo and nuclear endosperm are formed. At the seed-filling stage, other ovules have collapsed and the seed gradually comes to occupy the total volume of the pod. The fruit-to-seed length ratio decreases considerably during seed ripening. At fertilization, ovary length is four times greater than ovule length. In the mature state, the fruit and seed lengths are approximately equal. Seed size and weight diminish with an increase in seed number within a pod, although pod size remains constant. It is assumed that nonrandom abortion of young seeds inM. officinalis is under maternal control and is not related to structural abnormalities in ovule development or with limitation in pollen. We suppose that evolution of this species may have proceeded in the direction of a decrease in seed number and an increase in its sizes, which may play an important role in seed dispersal and seedling establishment.     

Does successful ovule development depend on its position within the pod? Examples from Neotropical Fabaceae

Previous studies have shown a nonrandom pattern of ovule fate probabilities according to ovule position in legume pods. Here, we tested how ovule position within the pods of two Fabaceae affects its fate. We expected higher proportion of well‐formed seeds near the fruit tips and of unfertilized and aborted ovules near fruit bases. We collected pods of Poincianella pyramidalis and Anandenanthera colubrina in a seasonal dry forest in northeastern Brazil and recorded total pod length and ovule number, position, and fate (unfertilized, well formed, aborted, and predated). The proportion of well‐formed ovules at fruit tips was significantly higher than at fruit bases in P. pyramidalis. The opposite pattern was found for unfertilized and aborted ovules, thus corroborating our hypothesis. However, the probability of seed predation in A. colubrina was significantly higher in pod tips, thus providing moderate support for our hypotheses. Interspecific differences in the patterns of ovule fate are likely to be driven by species pollination systems.     

Turgor-sensitive sucrose and amino acid transport into developing seeds of Pisum sativum. Effect of a high sucrose or mannitol concentration in experiments with empty ovules

Sugar and amino acid transport into empty ovules of Pisum sativum L. cv. Marzia was examined. In fruits containing 4–6 developing seeds, the embryo was removed from four ovules. After this surgical treatment, each empty seed coat was filled with a solution (pH 5.5) containing a low (0, 50 or 200 m M ), medium (350, 400 or 500 m M ) or high (0.7 or 1 M ) concentration of sucrose and/or mannitol. In pulse-labelling experiments with sucrose and α-aminoisobutyric acid (AIB), transport of sucrose and AIB into an empty ovule filled with a solution containing a high sucrose concentration was the same as transport into an ovule filled with a mannitol solution of similar osmolarity, demonstrating that a high sucrose concentration in the seed coat apoplast affects phloem transport of sucrose and AIB into the seed coat only by the osmotic effect. The osmolarity of a given solution filling the seed coat cavity appeared to be important for phloem transport of sucrose and AIB into empty ovules.
In our experiments, 350 m M appeared to be the optimal concentration for sucrose and AIB transport into the cavity within an empty ovule, giving results comparable with transport into intact ovules. A lower osmolarity of the solution induced less transport. Very high sucrose or mannitol concentrations caused a strong inhibition of sucrose and AIB unloading from the seed coat, so that transport into the empty ovules was inhibited. A low (strongly negative) but not too low osmotic potential of the solution in the seed coat apoplast seems necessary to maintain a normal rate of phloem transport into developing seeds. Apparently, the "sink strength" of developing seeds is turgor-sensitive.     

Pollination of Australian Macrozamia cycads (Zamiaceae): effectiveness and behavior of specialist vectors in a dependent mutualism

Complementary field and laboratory tests confirmed and quantified the pollination abilities of Tranes sp. weevils and Cycadothrips chadwicki thrips, specialist insects of their respective cycad hosts, Macrozamia machinii and M. lucida. No agamospermous seeds were produced when both wind and insects were excluded from female cones; and the exclusion of wind-vectored pollen alone did not eliminate seed set, because insects were able to reach the cone. Based on enclosure pollination tests, each weevil pollinates an average 26.2 ovules per cone and each thrips 2.4 ovules per cone. These pollinators visited similar numbers of ovules per cone in fluorescent dye tests that traced insect movement through cones. Fluorescent dye granules deposited by Cycadothrips were concentrated around the micropyle of each visited ovule, the site of pollen droplet release, where pollen must be deposited to achieve pollination. In contrast, Tranes weevils left dye scattered on different areas of each visited ovule, indicating that chance plays a greater role in this system. Each weevil and 25 thrips delivered 6.2 and 5.2 pollen grains, respectively, on average, to each visited ovule per cone, based on examination of dissected pollen canals. In sum, the pollination potential of 25 Cycadothrips approximates that of one Tranes weevil.     

Sex and pollen: the role of males in stabilising a plant-seed eater pollinating mutualism

Some plants are exclusively pollinated by an insect whose larvae feed on their seeds. The net outcome of a single visit for the plant depends on the number of ovules fertilised by the visitor, the number of eggs laid, and the number of seeds eaten by each larva. Unlike other known plant-seed eater pollinating mutualisms, the globeflower-globeflower fly mutualism (Trollius europaeus-Chiastocheta spp.) is unique in that not only females but also males visit flowers, and both sexes are potential pollinators. I analysed the relative efficiency of Chiastocheta males versus females in transporting pollen and fertilising globeflower ovules. I show that there is no sex-specific morphological adaptation or behaviour to enhance pollen collection and transportation in Chiastocheta flies, and that males contribute to pollination. However, because of their smaller body size, males transport significantly less pollen than females. Less seeds are produced after a visit from a male than after a visit from a female. A single female visit contributes to about 12% of total seed production, and a single male visit to only 5.4%. Females tend to spend more time inside the flower than males, and the number of ovules fertilised is significantly correlated with the time insects spent inside the closed corolla. The lower efficiency of ovule fertilisation by a male's single visit is compensated for by the higher rate of flower visitation by males: a flower receives about twice as many visits from males as from females during a time unit. The contribution of males to pollination is of major importance with respect to understanding the evolutionary stability of the globeflower-globeflower fly mutualism, as males satiate pollen requirement of flowers, masking the antagonistic effect of ovipositing females.     

Postfertilization causes of differential success of pollen donors in Erythronium grandiflorum (Liliaceae): nonrandom ovule abortion

Seed paternity in Erythronium grandiflorum does not fully reflect the proportion of pollen on the stigma. When two types of pollen are simultaneously applied to the stigma, outbred seeds are produced over inbred, and seeds from more distant donors are produced over seeds from donors nearby. I looked for postfertilization causes of these previously reported patterns of differential success of pollen donors. I simultaneously pollinated stigmas with pollen from two donors and observed ovule development through a window sliced in the ovary. Pollen donor pairs were self and cross, donors 1 and 100 m from the recipient, and two donors each 100 m from the recipient. Since one donor was always the alternate homozygote from the recipient at the malate dehydrogenase locus, I could determine the paternity of developing seeds. When it appeared that ovules were aborting, I removed them and determined their paternity using starch gel electrophoresis. Ovules fertilized by self pollen were more likely to abort than ovules fertilized by cross pollen, and ovules fertilized by nearby donors were more likely to abort than ovules fertilized by distant donors. Ovules fertilized by donors 100 m from the recipient were equally likely to abort. There was not a significant relationship between the proportion of ovules fertilized by a pollen donor and the probability of those fertilized ovules developing into seeds. There was no relationship between ovule position within a fruit and ovule abortion. I manipulated available resources by removing leaves and by permitting only one fruit to develop per plant. Decreasing the amount of resources increased the proportion of aborted ovules. Abortion of ovules of lesser quality appears to release resources that can then be used to develop other offspring.     

Rescuing Robert Brown—The Origins of Angio-Ovuly in Seed Cones of Conifers

Robert Brown (1827) recognized the distinction between the two major classes of seed plants on the basis, respectively, of exposed ovules, which receive pollen directly, and enclosed ovules, which do not, at the time of pollination. The two groups, unfortunately, became known as “Gymnosperms” and “Angiosperms”, a distinction not made by Brown. The names are, at best, useful only as identifying labels. As is established here, gymno-ovuly and angio-ovuly would have been better words and should serve as the most consistent distinguishing character among groups of seed plants. Although conifers can hardly be claimed to be members of any lineage that led to the Angiosperms, the gymno-ovulate method of reproduction must have occurred in the direct ancestors of flowering plants. Interpolating development and functional evidence serves to resurrect Robert Brown’s important but neglected observation and correct misstatements implied in generally accepted systematic terminology. The result of this approach is to discover that the conifers in their reproductive diversity show some trends indicative of those characters that define the flowering plants. Although non-homologous these character manifestations provide useful models which can lead to a better understanding of the origin of “Angiosperms” from “Gymnosperms.” In the complex and extended process of reproductive development in both major groups regular transpositions occur, depending on whether “gymno-”and “angio-”are applied to ovules or seeds. Gymno-ovuly and angio-spermy are the most common conditions in conifers but result in intrinsically opposed processes of pollen reception and subsequent seed protection, which are resolved by successive changes in seed-cone structure. The modern conifers illustrate considerable diversity of ovulate and seed cone structure in relation to several biological functions, including traits that anticipate the angio-ovulate condition, in cone inception, ovule enclosure, pollen and embryo development and seed dispersal. From the paleobotanical record we do have a good grasp of the structural changes in the progressive modification of the coniferous seed cone in conifers, starting with the antecedent Carboniferous Cordaitales. These changes have involved the progressive condensation of a lax, branched strobilus and end with the highly condensed uniovulate “cone” as in many Podocarpus species in which the equivalent of a bitegmic ovule exists. The progressive changes can be perceived typologically among existing conifers but need to be understood as the result of biological processes that suggest increased reproductive efficiency, and can be added to our present quite robust understanding of conifer phylogeny.     

POLLINATION,FERTILIZATION AND OVULE ABORTION IN OXALIS MAGNIFICA

Pollination, fertilization and ovule abortion were studied in Oxalis magnifica (Rose) Knuth, a strongly self-incompatible herb that regularly matures only a fraction of its ovules. Examination of cleared ovules indicated that among 9 individuals the average number of ovules fertilized ranged from 48–92%. The remaining ovules either failed to produce female gametophytes, or more commonly contained unfertilized female gametophytes, despite large numbers of compatible pollen grains that were placed on stigmas. Abortion of fertlized ovules could be detected first by the flattened and enlarged appearance of the endosperm nuclei, followed by visible deterioration of the embryo. Among individuals the rate of embryo abortion varied from 3.4–47.9%. At lower levels of pollination an almost one-to-one relationship existed between the number of pollen grains placed on stigmas and the number of seeds matured in the capsule. No threshold number of pollen grains necessary for successful pollen tube growth and fertilization could be demonstrated. Reduction in seed number through embryo abortion provides an opportunity for selection among developing seeds. The potential for this form of selection varies widely among individuals of Oxalis magnifica, which showed a 14-fold variation in the average percentage of aborted ovules.     

Hypotonic buffer induces meiosis and formation of anucleate cytoplasmic islands in the egg of the two-spotted cricket Gryllus bimaculatus

In insects, egg activation is known to occur in vivo and independently of fertilization, but its mechanisms are poorly understood. To gain understanding of these mechanisms, an attempt was made to activate the egg of Gryllus bimaculatus in vitro. It was found that meiosis resumed and was completed in unfertilized eggs treated with hypotonic buffer. Early developmental processes in activated, unfertilized eggs were investigated and compared with those in fertilized eggs. Mitosis did not progress, resulting in formation of anucleate cytoplasmic islands (pseudoenergids). Development in the activated, unfertilized eggs stopped at this stage and both yolk subdivision and cellularization did not occur. To elucidate the role of the nucleus in the developmental process to the syncytial stage in fertilized eggs, eggs were treated with aphidicolin to inhibit DNA polymerization. It was found that pseudoenergids also formed in these aphidicolin-treated fertilized eggs. These results demonstrate that pseudoenergids can increase in number independently of nuclei, suggesting that the cytoplasm rather than the nucleus plays the primary role in development to the syncytial stage in G. bimaculatus.     

Integration and macroevolutionary patterns in the pollination biology of conifers

Integration influences patterns of trait evolution, but the relationship between these patterns and the degree of trait integration is not well understood. To explore this further, we study a specialized pollination mechanism in conifers whose traits are linked through function but not development. This mechanism depends on interactions among three characters: pollen that is buoyant, ovules that face downward at pollination, and the production of a liquid droplet that buoyant grains float through to enter the ovule. We use a well‐sampled phylogeny of conifers to test correlated evolution among these characters and specific sequences of character change. Using likelihood models of character evolution, we find that pollen morphology and ovule characters evolve in a concerted manner, where the flotation mechanism breaks down irreversibly following changes in orientation or drop production. The breakdown of this functional constraint, which may be facilitated by the lack of developmental integration among the constituent traits, is associated with increased trait variation and more diverse pollination strategies. Although this functional “release” increases diversity in some ways, the irreversible way in which the flotation mechanism is lost may eventually result in its complete disappearance from seed plant reproductive biology.     

Development of fertilized ovules and their role in the growth of the pea pod

The histological development of fertilized ovules during fruit-set and development in pea ( Pisum sativum L. cv. Alaska) has been investigated. Killing the ovules on day 0 (anthesis) or day 1 prevented fruit-set and resulted in ovary degeneration. When the ovules were destroyed at later stages the ovaries developed, though the rate of growth of the pod was reduced significantly. Pollination in pea occurs normally the day before anthesis, and fertilization of the egg cell 32 to 48 h later. The first divisions of the zygote and endosperm nuclei started simultaneously (ca 48 h after pollination) but the endosperm developed more rapidly than the embryo; the embryo sac cavity was lined with free endosperm nuclei at the time of beginning suspensor elongation. Extracts of endosperm and ovule coats from ovules at day 7 after anthesis showed fruit-set activity in pea, the latter material having about 3 times more activity than the former per ovule basis. These results indicate that fertilization of the ovule is necessary for fruit-set in pea, and that compounds which induce fruit-set are probably synthesized in the ovules following fertilization.     

Resolving the systematic and phylogenetic position of isolated ovules: a case study on a new genus from the Permian of China

Isolated ovules occur in many fossil plant assemblages, where they provide important insights into seed‐plant diversity and evolution. However, in many cases, the ovules cannot be attributed to individual groups of seed plants, restricting systematic and evolutionary assessments that can be made from otherwise well‐characterized fossil taxa. In the present paper, we describe a new kind of ovule discovered in tuffaceous sediments from the Permian‐aged Xuanwei Formation in Guizhou Province, China. This ovule has 180° rotational symmetry and an integument comprising a variably thick sarcotesta, a uniformly thick sclerotesta and a uniformly thin endotesta. The nucellus is attached to the integument at least basally and contains a collapsed seed megaspore; a nucellar apex is absent. Both the integument and nucellus are vascularized by paired bundles in the major plane of the ovule; the integumentary bundles are considerably larger than the nucellar bundles and the nucellar bundles emerge from a conical vascular pad. Generation of a three‐dimensional reconstruction based on serial peels revealed the gross morphology and organization of the ovule and highlighted the presence of features consistent with cardiocarpalean‐type ovules (ovule shape, histological features of the integument) and also features more typical of lagenostomalean‐ and trigonocarpalean‐type ovules (large integumentary bundles, presence of nucellar bundles). To assess the affinity and evolutionary significance of the ovule, it has been included in a cladistic matrix of cardiocarpalean‐, lagenostomalean‐ and trigonocarpalean‐type ovules. Results place the ovule within the cardiocarpalean group of ovules known to have been produced by several plant groups, including cordaitean coniferophytes, pteridosperms and Palaeozoic conifers. The cladistic topology supports generic level distinction of the present species, requiring the establishment of Muricosperma guizhouensis Seyfullah & J.Hilton gen. & sp. nov . Lagenostomalean ovules produced by hydrasperman pteridosperms form a basal paraphyletic grade, whereas trigonocarpalean ovules produced by medullosan pteridosperms form a monophyletic group in which Stephanospermum is paraphyletic with respect to Rhynchosperma and Pachytesta. The results also place the Mississippian ovule Mitrospermum bulbosum apart from all of the Pennsylvanian species of Mitrospermum that form a strongly supported clade. Consequently, M. bulbosum is transferred to the new genus Whitaddera Seyfullah & J.Hilton as W. bulbosa (Long) Seyfullah & J.Hilton. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 164 , 84–108.     

Self pollination and resource availability affect ovule abortion inCassia fasciculata (Caesalpiniaceae)

We examined the effects of pollen source and resource availability on ovule abortion in the annual legumeCassia fasciculata. Pollen source was controlled by hand-pollinating flowers with cross- or self-pollen. Resource availability to developing fruits was controlled by adjusting fruit loads (heavy versus light) on each plant and exposing plants to different photoperiod cycles (16 vs 12 h of light; short days favor fruit growth at the expense of vegetative growth). In mature fruits the proportion of ovules expanding (showing some development over virgin ovules) ranged from 89–95% and did not increase with resource availability, suggesting that unexpanded ovules were either unfertilized or obligately aborted shortly after fertilization. The proportion of expanded ovules maturing in mature fruits was near 97% for both self- and crosspollinations in the treatment with highest resource availability (light load, short days) and lower in the remaining treatments, where self-pollination resulted in up to 9% lower seed maturation than cross-pollination. In the latter three treatments a pollen source effect was dependent upon the maternal plant; in some plants selfing increased abortion and in others it did not. Collectively, the results suggest that (1) both pollen source and resource availability affect ovule abortion, (2) at least some abortion is facultative, and (3) when resources are limited, self-pollination increases abortion in some but not all maternal plants.     

Reciprocal inheritance of centrosomes in the parthenogenetic hymenopteran Nasonia vitripennis

Background: In the majority of animals, the centrosome-the microtubule-organizing center of the cell-is assembled from components of both the sperm and the egg. How the males of the insect order Hymenoptera acquire centrosomes is a mystery, as they originate from virgin birth.Results: To address this issue, we observed centrosome, spindle and nuclear behavior in real time during early development in the parthenogenetic hymenopteran Nasonia vitripennis. Female meiosis was identical in unfertilized eggs. Centrosomes were assembled before the first mitotic division but were inherited differently in unfertilized and fertilized eggs. In both, large numbers of asters appeared at the cortex of the egg after completion of meiosis. In unfertilized eggs, the asters migrated inwards and two of them became stably associated with the female pronucleus and the remaining cytoplasmic asters rapidly disappeared. In fertilized eggs, the Nasonia sperm brought in paternally derived centrosomes, similar to Drosophila melanogaster. At pronuclear fusion, the diploid zygotic nucleus was associated only with paternally derived centrosomes. None of the cytoplasmic asters associated with the zygotic nucleus and, as in unfertilized eggs, they rapidly degenerated.Conclusions: Selection and migration of the female pronucleus is independent of the sperm and its aster. Unfertilized male eggs inherit maternal centrosomes whereas fertilized female eggs inherit paternal centrosomes. This is the first system described in which centrosomes are reciprocally inherited. The results suggest the existence of a previously undescribed mechanism for regulating centrosome number in the early embryo.