Pistil traits and flower fate in apricot (Prunus armeniaca)

Although pollination is essential for both seed and fruit set in most angiosperms, even after an adequate pollination, only a fraction of the flowers develop into fruits. The role played by floral traits on reproductive success is well known, but the possible influence of pistil traits has been overlooked, probably because of the difficulty of non-destructive pistil examination. The aim of this work was to examine the influence of several pistil traits on reproductive success in apricot ( Prunus armeniaca ). For this purpose, in a population of individually labelled flowers, the styles were cut off once the pollen tubes had reached the ovary but prior to the achievement of fertilisation. This approach allowed relating several morphological and physiological pistil parameters in the dissected styles and stigmas to the subsequent set or abscission of the corresponding ovary that remained in the plant. Under the same pollination conditions, the flowers that finally set a fruit show a larger stigmatic area and a higher number of pollen grains, pollen tubes growing along the style and xylem vessels surrounding the transmitting tissue than flowers that abscise before the establishment of fruit set. Furthermore, starch is present in the transmitting tissue of the style in all the flowers that develop into fruits but only in half of the flowers that abscise. The examined pistil traits established prior to fertilisation are related to flower fate, suggesting that the capacity of a flower to become a fruit could be preconditioned at anthesis.     

Ovary starch reserves and pistil development in avocado (Persea americana)

In avocado, only a very small fraction of the flowers are able to set fruit. Previous work in other woody perennial plant species has shown the importance of carbohydrates accumulated in the flower in the reproductive process. Thus, in order to explore the implications of the nutritive status of the flower in the reproductive process in avocado, the starch content in the pistil has been examined in individual pollinated and non‐pollinated flowers at anthesis and during the days following anthesis. Starch content in different pistilar tissues in each flower was quantified with the help of an image analysis system attached to a microscope. Flowers at anthesis were rich in highly compartmentalized starch. Although no external morphological differences could be observed among flowers, the starch content varied widely at flower opening. Starch content in the ovary is largely independent of flower size because these differences were not correlated with ovary size. Differences in the progress of starch accumulation within the ovule integuments between pollinated and non‐pollinated flowers occurred concomitantly with the triggering of the progamic phase. The results suggest that starch reserves in the ovary could play a significant role in the reproductive process in avocado.     

Self‐pollination and parthenocarpic ability in developing ovaries of self‐incompatible Clementine mandarins (Citrus clementina)

This study aimed to determine if self‐pollination is needed to trigger facultative parthenocarpy in self‐incompatible Clementine mandarins (Citrus clementina Hort. ex Tan.). ‘Marisol’ and ‘Clemenules’ mandarins were selected, and self‐pollinated and un‐pollinated flowers from both cultivars were used for comparison. These mandarins are always seedless after self‐pollination and show high and low ability to develop substantial parthenocarpic fruits, respectively. The time‐course for pollen grain germination, tube growth and ovule abortion was analyzed as well as that for carbohydrates, active gibberellins (GA₁ and GA₄), auxin (IAA) and abscisic acid (ABA) content in the ovary. ‘Clemenules’ showed higher pollen grain germination, but pollen tube development was arrested in the upper style 9 days after pollination in both cultivars. Self‐pollination did not stimulate parthenocarpy, whereas both un‐pollinated and self‐pollinated ovaries set fruit regardless of the cultivar. On the other hand, ‘Marisol’ un‐pollinated flowers showed greater parthenocarpic ovary growth than ‘Clemenules’ un‐pollinated flowers, i.e. higher ovule abortion rate (+21%), higher fruit set (+44%) and higher fruit weight (+50%). Further, the greater parthenocarpic ability of ‘Marisol’ paralleled higher levels of GA₁ in the ovary (+34% at anthesis). ‘Marisol’ ovary also showed higher hexoses and starch mobilization, but lower ABA levels (?64% at anthesis). Self‐pollination did not modify carbohydrates or GA content in the ovary compared to un‐pollination. Results indicate that parthenocarpy in the Clementine mandarin is pollination‐independent with its ability to set depending on the ovary hormone levels. These findings suggest that parthenocarpy in fertile self‐incompatible mandarins is constitutively regulated.     

The roles of abscisic acid and ethylene in the abscission and senescence of cocoa flowers

Cocoa flowers have a limited period of longevity; more than 90% of unpollinated flowers abscised within 32 h after anthesis. Abscisic acid (ABA) levels increased significantly prior to abscission. By 21 h after anthesis, ABA levels had increased almost 10-fold, and by 32 h flowers had 20-fold higher levels of ABA than at anthesis. Fluridone completely inhibited both the increase in ABA, the formation of an abscission zone, and the abscission and senescence of flowers. In contrast, ethylene production increased only slightly 21 h after anthesis and was only 2-fold higher after 32 h. Aminoethoxyvinylglycine (AVG) delayed but did not prevent abscission. In cocoa flowers, ABA is the primary regulator of abscission; ethylene accelerates abscission but only in the presence of ABA. Naphthalene acetic acid (NAA) treatment of flowers at anthesis prevented abscission zone formation and flower abscission, but did not induce fruit set. All parts of the NAA-treated flower except the pedicel senesced after 6 days. NAA+AVG treatment only delayed, whereas fluridone treatment completely prevented flower senescence.     

Flower abscission and inflorescence carbohydrates in sensitive and non-sensitive cultivars of grapevine

The Gewurztraminer (GW) and the Pinot noir (PN) cultivars of grapevine differ in their sensitivity to environmental factors that can cause flower abscission, cv. GW being the most sensitive. In order to further define the mechanisms leading to abscission, and owing to the importance of sugars in the achievement of sexual organ ontogenesis, we attempted to correlate the chronology of flower ontogenesis with the variations of carbohydrates in the inflorescence. In the vineyard, under optimal climatic conditions, fruit set of cv. GW and cv. PN was 82% and 65%, respectively. The sugar distribution was different in their inflorescences during the entire duration of flower development. Between stages 15 and 17, flowers of GW and PN reached the crucial meiosis stage. At that time, the inflorescences of cv. GW exhibited higher concentrations of starch and sucrose, whereas those of PN presented higher levels of glucose and fructose. Despite higher starch concentrations in GW inflorescences, starch reserves were present in the ovules and anthers of PN but not in those of GW. These results suggest that the higher content of reserve and transport carbohydrates in the inflorescences of GW favour flower development and fruit set under optimal environmental conditions. Furthermore, since meiosis represents a key step of female development, the different sugar concentrations in the inflorescences of the two cultivars at stages 15 and 17 could be related to the sensitivity to flower abscission under climatic stress. In particular, the presence of starch granules in PN ovules and anthers might explain the higher resistance of this cultivar to flower abscission.     

Ovary starch reserves and flower development in apricot (Prunus armeniaca)

In histerant species where flowering takes place prior to leaf emergence, a flower lifespan occurs in the absence of new photoassimilates and at the expense of pre-stored reserves either in the plant as a whole or in the flower itself. In the present study, the role that the photoassimilates stored in the flowers might play in flower development from anthesis to fertilization in Prunus armeniaca L. (apricot), a histerant species, was explored. Starch content in individual flowers was measured with the help of an image analysis system. Starch content decreased from its highest value at anthesis and disappeared from the ovary 9 days later. This decrease was inversely related to an increase in ovary size and in cell number in the pericarp, suggesting an intraflower, self-supported development. This process is conserved in both pollinated and nonpollinated flowers and therefore seems to be inherent to the flower at anthesis. The onset of fruiting is preceded by the establishment of large differences among ovaries; while some experience continuous growth, others stop growing and eventually drop. Interestingly, large differences in starch content are found among flowers at anthesis. These results are discussed in terms of the possible implications of pre-stored starch in the flower supporting initial flower development.     

Effect of ovular receptivity on seed set and fruit development in cucumber (Cucumis sativus L.)

Summary Seed set and fruit development in cucumber (Cucumis sativus L.) were studied in relation to female flower receptivity from day — 2 before anthesis to day + 2 after anthesis. The female cucumber flower is protogynous. The pistil was receptive 2 days before anthesis. The iso-electric focusing (IEF) patterns of the stigma/style proteins, were identical from day -5 to day +2. In pollinated flowers in vivo germination and pollen-tube growth in the ovary were affected by pistil age from day -2 to day +2. In addition, differences in sectorial filling in full seeds were observed within the fruits. A negative correlation was observed between the frequency of fertilized ovules in the pedoncular part of the fruit and ovary length at the time of pollination. In the whole fruit, significant differences in the number of full seeds and fruit size at maturity were found, and these were observed to be correlated with the various stages of female flower maturation at pollination. The day -2 and day +2 stages yielded the smallest fruits with few full seeds compared to the day -1, day 0 and day +1 stages, which had the biggest fruits and a large number of full seeds. A strong positive correlation was found between total seed number (including full and empty seeds), fruit length and weight at maturity. All these results suggest that both seed set in the different parts of the fruit and fruit development are controlled by ovular receptivity rather than by stigma/style receptivity.     

Changes of Carbohydrates in Pepper (Capsicum annuum L.) Flowers in Relation to Their Abscission Under Different Shading Regimes

Abscission of pepper flowers is enhanced under conditions oflow light and high temperature. Our study shows that pepperflowers accumulate assimilates, particularly in the ovary, duringthe day time, and accumulate starch, which is then metabolizedin the subsequent dark period. With the exception of the petals,the ovary contains the highest total amounts of sugars and starch,compared with other flower parts and contains the highest totalactivity, as well as activity calculated on fresh mass basis,of sucrose synthase, in accordance with the role of this enzymein starch biosynthesis. Low light intensity or leaf removaldecreased sugar accumulation in the flower and subsequentlycaused flower abscission. The threshold of light intensity fordaily sugar accumulation in the sink leaves was much lower thanin flowers, resulting in higher daytime accumulation of sugarsin the sink leaves than in the adjacent flower buds under anylight intensity, suggesting a competition for assimilates betweenthese organs. Flowers of bell pepper cv. ‘Maor’and ‘899’ (sensitive to abscission) accumulatedless soluble sugars and starch under shade than the flowersof bell pepper cv. ‘Mazurka’ and of paprika cv.‘Lehava’ (less sensitive). The results suggest thatthe flower capacity to accumulate sugars and starch during theday is an important factor in determining flower retention andfruit set. Pepper; Capsicum annuum L.; abscission; shading; pepper flowers; ovary; leaves; sugars; starch; acid invertase; sucrose synthase     

SECONDARY-FRUIT ONTOGENY IN NAVEL ORANGE

Ontogeny of the secondary fruit of navel orange (Citrus sinensis [L.] Osbeck) was studied using flower buds and fruit from groves in north central Florida. The secondary fruit, or navel, develops as a whorl of secondary-carpel primordia within the primary-carpel whorl when the flower bud is 1.5 to 2.0 mm in length. A complete secondary gynoecium with fused ovary but separate styles and stigmas develops within the primary gynoecium before anthesis. The stigma and style of secondary carpels are not as distinct as those of primary carpels. Three types of tissue protrusions from the secondary fruit into primary-fruit locules were abnormal placentae, free secondary carpels, and secondary-carpel outgrowths, the last being the most common. An abscission layer is present in the central axis of the primary fruit near the base of the secondary ovary. This layer is indistinguishable from adjacent tissues prior to commencement of secondary fruit abscission. Parenchyma cells in the abscission layer, however, contain large numbers of starch grains during abscission. Secondary and primary fruit have similar sigmoid growth patterns; however, onset of the cell-enlargement stage is approximately 2 wk later in secondary fruit.     

Carbohydrate Availability in Relation to Fruitlet Abscission in Citrus

Abscission of flowers and fruitlets in the Washington navelorange (Citrus sinensis [L.] Osbeck) has been characterizedin relation to carbohydrate availability. A main wave of flowerabscission occurs shortly after anthesis while the carbohydratereserves in the tree are high. Fruitlet abscission starts approx.30 d after the commencement of flowering, while carbohydrates(mainly starch) are being accumulated in the leaves. Flowerand early fruitlet abscission are not caused by carbohydrateshortage. During late fruitlet abscission sucrose concentrationin the leaves falls to a low value demonstrating a limitationin supply and competition among the developing fruitlets forcarbohydrates. Concentrations of sucrose and reducing sugarsin the peel of the fruitlets also fall to low values, and arelationship could be demonstrated between these free sugarlevels and abscission. Ringing increases carbohydrate supplyto fruit and reduces late fruitlet abscission, but only hasa marginal effect on the growth of the fruitlets, which seemsless sensitive than abscission to carbohydrate shortage. Thelimitation of carbohydrate supply to the fruitlets occurs whilestarch levels in the leaves remain high. Slow mobilization ofstarch reserves may be one factor limiting set in Citrus. Copyright2001 Annals of Botany Company Carbohydrate supply, citrus, fruit growth and abscission, ringing, navel orange, starch, sugar metabolism     

Biochemical characterization of soybean ovary growth from anthesis to abscission of aborting ovaries

Soybean (Glycine max [L.] Merr.) ovary growth was measured from anthesis to 6 days after anthesis (DAA) to establish a timetable of biochemical events that might be useful in identifying processes that initiate abscission. Two procedures were developed to provide samples with either high or low percent pod set for `IX93-100,' a semideterminate line having long racemes. Characteristics measured were fresh and dry weight, soluble and insoluble protein, soluble carbohydrate, starch, RNA, and DNA. Setting ovaries grew more rapidly than abscising ovaries. Since there was a daily increase in ovary weight in both groups, all measured characteristics showed daily increases when expressed on perovary basis. Statistically significant differences between groups were detected between 2 and 5 DAA for most characteristics. When chemical composition was expressed on concentration basis, starch level was significantly higher in setting ovaries at 5 and 6 DAA. Regression analysis showed that these deviations between setting and abscising samples started between anthesis and 1 DAA. We conclude that processes leading to eventual shedding of fertilized ovaries (called flower abortion in soybeans) commence soon after anthesis of the shed flower, and that setting and abscising ovaries do not differ in protein, soluble carbohydrate, starch, or nucleic acid content when abscission processes begin.     

Flowering and Fruiting Phenology of Magnolia hypoleuca

Abstract The floral biology of Magnolia hypoleuca , a tall tree species widely distributed in Japan, was studied in a deciduous broad-leaved forest in Hokkaido near the northern limit of distribution of the species. The flower is large, protogynous and nectarless. Its petals close after the female stage and again after the male stage. The mean duration of flowering of an individual flower is 3–4 days, while the flowering period of an individual tree lasts up to 40 days. The main pollinators of the flowers were beetles. Success of pollination is assumed to be affected by weather conditions at anthesis. Average fruit sets (ratios of fruits to flowers) were 13–25%. Hand-pollination of 44 flowers produced a significantly higher fruit set (28 fruits) than control flowers (33/133). These results suggest that the fruit set is limited by pollination. The frequency of infertile ovules was higher in controls than in hand-pollinated fruits, indicating that the seed set was also limited by pollination. We propose a mimicry hypothesis in which the non-rewarding female-stage flower mimics the rewarding male-stage flower to explain the short life-span of an individual flower, the long flowering period of an individual tree, and the coexistence of flowers of various stages on a single tree.     

The Significance of the Obturator in the Control of Pollen Tube Entry into the Ovary in Peach (Prunus persica)

Changes in the obturator of the peach (Prunus persica) havebeen investigated and related to pollen tube growth in thisregion. At anthesis, the cells of the obturator are active andrich in starch reserves. Twelve days after anthesis these cellsproduce a secretion that stains for carbohydrates and for proteins.As the secretion is produced, starch vanishes from these cellsand they degenerate and collapse as callose is accumulated.Secretion is independent of pollination as it takes place ina similar fashion both in pollinated and in unpollinated flowers. Pollen tube growth along the obturator surface depends on thissecretion for, although pollen tubes reach the base of the styleseven days after pollination, they cannot grow on the obturatoruntil five days later, when the secretion is produced. Thisdiscontinuous secretion taking place at the obturator may providea mechanism that controls the entrance of pollen tubes intothe ovary in the peach. Prunus persica, peach, obturator, pollen tube     

Morphological and cytological development and starch accumulation in hermaphrodite and staminate flowers of olive (Olea europaea L.)

In olive (Olea europaea L.), the formation of functionally staminate flowers rather than fully functional hermaphrodites is one of the major factors limiting fruit set, as flowers with aborted pistils are incapable of producing fruit. Studies conducted on various angiosperm species have shown a correlation between flower abortion and starch content. Thus, it is important to know if starch content plays a role in regulating pistil development in olive and if so, what mechanism regulates starch distribution. Cyto-histological observations of staminate and hermaphrodite olive flowers show that pistil development in staminate flowers is interrupted after the differentiation of the megaspore mother cell. At that stage, starch grains were only detected in the ovary, style and stigma of the hermaphrodite flowers. No starch was observed in the pistils of the staminate flowers. This finding suggests a tight correlation between starch content and pistil development. The secondary origin of starch within the flower is indicated by low chlorophyll content in the gynoecium, undetectable Rubisco activity in the pistils of these two kinds of flowers and by the ultrastructure of the plastids observed by transmission electron microscope analysis. The plastids have few thylakoid membranes and grana and in the staminate flowers appeared very similar to proplastids. Considering differences in starch content between staminate and hermaphrodite flowers and the secondary origin of the starch, differences in pistil development in the staminate and hermaphrodite flowers could be related to differences in the sink strength of these two types of flowers.     

Studies on flower longevity in Digitalis

The flowers of Digitalis purpurea respond to pollination by rapid corolla abscission without any loss of corolla turgor, nor any significant loss of corolla constituents, relative to the corollas of unpollinated flowers of a similar age. The corollas of unpollinated flowers too eventually abscise, 6 d after the stigma opens, however, they do so with only a minimal loss of fresh weight or corolla constituents. Pollination causes an increase in ethylene production detectable within 1 h. Increased ethylene production occurs initially only from the upper portion of the style, later from the lower portion, and lastly, between 23 and 48 h after pollination, from the ovary plus calyx. The pollination response can be induced by exogenous ethylene, the degree of weakening of the corolla abscission zone being dependent upon the concentration and duration of the exposure period and on the stage of flower development. The regulation of ethylene biosynthesis and its involvement in the control of pollination-induced corolla abscission are discussed.     

Studies on flower longevity in Digitalis

Flower lifespan was terminated by corolla abscission 5–6 days after stigma opening in the unpollinated flower. Increased pollen loads produced increased seed set and reduced flower longevity progressively to a minimum of one day after pollination with pure pollen. Weakening of the abscission zone was detectable 8 h after pollination, whilst the pollen tubes were still within the stigmatic zone, suggesting that a stimulus, moving at 4 mm h^–1 minimum, was transmitted through the style and ovary. Soon after pollination removal of the stigma prevented the pollination-induced corolla abscission. Later it was necessary to remove the stigma and upper style, and later still the whole style to delay abscission. The progressive induction of the stigma and style took place at a rate of 1.5 mm h^–1, in advance of the pollen tubes which grew at 0.75 mm h^–1. It was not possible to reproduce the pollination effects by application of indoleacetic acid (IAA) to the stigma or the style.Abbreviation IAA Indoleacetic acid     

Floral biology and pollination in Cucumis melo L., a tropical andromonoecious cucurbit

We studied floral biology and the role of flower visitors in fruit set of muskmelon (Cucumis melo L.). Muskmelon produced Staminate (hereafter ‘S’) and Hermaphrodite (hereafter ‘H’) flowers with a ratio of 18:1. Flowers opened around 40 days after planting and flowering continued for another 40 days. Longevity of both types of flowers was one day. Anthesis in S flowers was earlier (0545–0700 h) than that of H flowers (0700–0830 h). Anther dehiscence occurred 10–15 min after anthesis in both the flowers. The number of pollen grains in H flowers was significantly lower (2715 ± 143) than S flowers (4968 ± 398). Pollen viability and stigma receptivity was maximum between 1600 and 1800 h. The pollen tube of pollen grains from H flowers grew very slowly and never reached the ovary. There were two peaks in nectar production in a day (1000–1200 h and 1400–1800 h). Sixteen species of insects belonging to Hymenoptera (81.25%), Diptera (12.50%) and Lepidoptera (6.5%) visited muskmelon flowers. Major visitors were honeybees, Apis cerana F. and A. florea F. and both were efficient pollinators. Diversity and abundance of flower visitors positively correlated with floral abundance. In pollination exclusion experiments, no fruit set was observed in bagged, self-pollinated and the flowers artificially crossed only with other H pollen. But, those flowers hand pollinated with pollen from S flowers and ones that were allowed for open pollination set maximum fruits. The crop needs a minimum of 8–10 bee visits per flower to achieve the same level of pollination efficiency provided with natural/unrestrained visits.     

Wind-dragged corolla enhances self-pollination: a new mechanism of delayed self-pollination

BACKGROUND AND AIMS: Delayed self-pollination is a mechanism that allows animal-pollinated plants to outcross while ensuring seed production in the absence of pollinators. This study aims to explore a new mechanism of delayed self-pollination facilitated by wind-driven corolla abscission in Incarvillea sinensis var. sinensis. METHODS: Floral morphology and development, and the process of delayed self-pollination were surveyed. Experiments dealing with pollinator and wind exclusion, pollination manipulations, and pollinator observations were conducted in the field. KEY RESULTS: Delayed self-pollination occurs when the abscising corolla driven by wind drags the adherent epipetalous stamens, thus leading to contact of anthers with stigma in late anthesis. There is no dichogamy and self-incompatibility in this species. The significantly higher proportion of abscised corolla under natural conditions as compared with that in wind-excluding tents indicates the importance of wind in corolla abscission. When pollinators were excluded, corolla abscission significantly increased the number of pollen grains deposited on the stigma and, as a result, the fruit and seed set. Only half of the flowers in plots were visited by pollinators, and the fruit set of emasculated flowers was significantly lower than that of untreated flowers in open pollination. This species has a sensitive stigma, and its two open stigmatic lobes closed soon after being touched by a pollinator, but always reopened if no or only little pollen was deposited. CONCLUSIONS: This delayed self-pollination, which involved the movement of floral parts, the active participation of the wind and sensitive stigma, is quite different from that reported previously. This mechanism provides reproductive assurance for this species. The sensitive stigma contributes to ensuring seed production and reducing the interference of selfing with outcrossing. The pollination pattern, which combines actions by bees with indirect participation by wind, is also a new addition to ambophily.     

Flower bud differentiation and development in fruiting and non-fruiting shoots in relation to fruit set in apricot (Prunus armeniaca L.)

Situations of high flower bud drop and low fruit set without apparent causes are common in fruit trees. The term flower quality has been coined to explain differences among flowers in their capacity to set fruit, but the causes underpinning these differences are largely unknown. This lack of knowledge is based on the fact that these differences are established a posteriori and there are no criteria to determine a priori what will make a flower to set a fruit or to drop. In this work, we profit from the empirical knowledge that there are fruiting and non-fruiting shoots to explore to which extent flower bud differentiation and bud development will affect the subsequent fruit set. For this purpose, the processes from flower bud differentiation to fruit set were sequentially analyzed in both types of shoots, over 2 years. More than half of the buds from long shoots aborted development and dropped before flowering. At anthesis, most of the remaining flowers showed underdeveloped pistils that failed to sustain pollen germination or pollen tube growth along the pistil. This unsuccessful development resulted in clear differences in fruit set between both types of branches. These results highlight that flower bud differentiation and development play an important role for fruit set and that developmental timing appears critical to reach anthesis with a fully developed pistil.     

Pollen-pistil interaction in kiwifruit (Actinidia deliciosa; Actinidiaceae)

The pollen-pistil interaction has been examined in kiwifruit (Actinidia deliciosa). In this species a large number of seeds are produced in each fruit and a great many pollen grains germinate and grow to reach the ovules. This growth is assisted by an abundant secretion that is present all along the pistilar tract. At anthesis, the stigma is covered by a secretion where the pollen grains germinate and grow. The stylar transmitting tissue is initially rich in starch reserves, but the starch gradually disappears and, concomitantly, an abundant secretion that stains for carbohydrates appears in all of the intercellular spaces. Pollen tube growth relies on this secretion since it is depleted after pollen tube passage, while in unpollinated flowers it remains unaltered throughout the flower life-span. In the ovary a similar situation occurs. The placental surface, where the pollen tubes grow before reaching the ovules, is covered by a number of obturators. At anthesis, these obturators are rich in starch reserves and have an abundant secretion on their outer surface. As time passes, starch disappears while the secretion increases. It is in this secretion that the pollen tubes grow on their way toward the ovules. These observations are discussed in terms of the support given by the pistil to pollen tube growth to achieve the highly successful reproductive performance of this species.