Intra‐specific pollen polymorphism in Mimosa pigra (Mimosaceae)

Five different types of pollen tetrads were detected from the short and long stamens of the male and hermaphrodite flowers of the androdioecious species, Mimosa pigra L. Light (LM) and scanning electron microscopes (SEM) were used in this study to redescribe the pollen forms and to identify the various pollen types encountered. An identification key and relative percentages of these pollen types were also presented. The study showed that the long stamens of the male flowers and those of the hermaphrodite ones have five (I, II, III, IV and V) and two (I and V) pollen types respectively, whereas the short stamens of both male and hermaphrodite flowers have only one pollen type (type I). We report here the first incidence of intra‐specific pollen polymorphism in androdioecious species.     

CLEISTOGAMY IN SALPIGLOSSIS SINUATA

Cleistogamy in Salpiglossis sinuata L. (Solanaceae) involves a series of developmental events that includes germination of pollen tetrads en masse inside the anther, penetration of pollen tubes through the anther wall toward the stigma which is in contact with the anther, growth of pollen tubes down the style, and, finally, fertilization. This occurs within 6 days from the first appearance of the flower as compared to 10–12 days for normal anthesis of chasmogamous flowers. Coincident with these events in cleistogamous flowers are lack of corolla development and reduced size of androecium and gynoecium.     

Dioecy and wind pollination in Pernettya rigida (Ericaceae) of the Juan Fernández Islands

Pernetttya rigida is endemic to the Juan Fernández archipelago. Although all flowers are complete, with seemingly fertile stamens and pistils, differences in fruit production and detailed field, anatomical, and morphological studies indicate they are functionally unisexual, and the species is consequently dioecious. A comparison of 15 features demonstrated sigdicant differences between the sexes. The populational sex ratio is 1:1. Nectaries located between the filaments produce small amounts of floral nectar with similar sugar composition in both sexes. There are =33,357 (or = 133,429) pollen tetrads (or pollen grains)/male flower and =109 ovules/female flower. No tetrads in either hand self-pollinated or open pollinated male flowers showed any pollen germination. Tetrads on open- and hand-pollinated female flowers germinate. Female flowers do not show autogamy or apomixis. During more than 80 hours of field observation, we recorded only seven floral visitors (representing three insect species). In spite of this, openpollinated female flowers have abundant fruit and seed-set. Thus, we conclude that pollen is transferred abiotically and the ever-present wind over the exposed ridges of the islands is the likely dispersal agent. A number of anemophilous features, such as dry pollen and exposed habitat, support this conclusion. Thus, dioecy and anemophily have evolved independently, in situ , in this species in this remote locality. Preservation of habitat and elimination of competitive invasives are the primary conservation challenges.     

Temporal patterns of seed set and decelerating fitness returns on female allocation in Zigadenus paniculatus (Liliaceae), an andromonoecious lily

I describe temporal patterns of seed production in the andromonoecious lily Zigadenus particulatus. Fruit set per flower and seed set per fruit declined through time within plants. Hand pollination experiments showed that this was not due to increasing pollen limitation. Nutrient supplementation had little effect on seed output, but leaf clipping reduced seed production, especially in late-blooming flowers, and removal of early-blooming flowers increased seed set by later flowers. Thus, the temporal pattern of seed output was due to declining availability of photosynthates. Plants with larger bulbs produced larger inflorescences, a greater proportion of hermaphrodite flowers, more fruits per hermaphrodite flower, and more seeds per fruit, but lost a greater fraction of their initial bulb mass as a consequence of fruiting. After controlling for the effects of bulb mass, plants with larger inflorescences produced a greater proportion of male flowers, and plants with more hermaphrodite flowers produced fewer fruits per hermaphrodite flower and fewer seeds per fruit. Thus, the female fitness gain curve was decelerating. The temporal decline in seed output provides a partial explanation for the parallel decline in allocation to pistils. However, a complete explanation for the pattern of gamete packaging requires an understanding of factors controlling male, as well as female, fitness.     

Population Size,Pollination and Breeding System of Silene stockenii Chater (Caryophyllaceae), an Annual Gynodioecious Species of Southern Spain

The distribution area, phenology, sex polymorphism, floral characteristics and breeding system of Silene stockenii (Caryophyllaceae), a narrowly endemic annual species of southern Spain, were studied. Only five populations were found in a total area of 2 ha. Silene stockenii is a gynodioecious species with fully female, fully hermaphrodite and intermediate plants bearing hermaphrodite, male-sterile, and partially male-sterile flowers. Male-sterile flowers are typically smaller than hermaphrodites. Nectar production was significantly higher in hermaphrodite plants and during the female phase of hermaphrodite flowers. The red flowers appear during the spring (March-May) and are pollinated by long-tongued Bombyliidae. Hand pollinations revealed that the species is self-compatible; however, natural self-pollination is rare due to marked protandry. Hand pollination significantly increased the number of seeds per fruit and seed set, indicating limited pollination in the field. In controlled pollinations female plants of S. stockenii produced higher seed set than hermaphrodite plants, but in freely pollinated plants fruit set and seed production was similar in both morphs, indicating that pollinators do not discriminate in favour of hermaphrodite plants.     

Pollen morphology ofLactoridaceae

The pollen wall ultrastructure of the primitive AngiospermLactoris fernandeziana Phil. (Lactoridaceae) is described. The monosulcate aperture, granular wall structure and sacci (all primitive features) suggest placement of this family in theMagnoliales. Pollen ofLactoris is compared to fossil dispersed pollen from the Lower and lower-Upper Cretaceous. The fossil pollen shares characteristics which are restricted to theLactoridaceae suggesting that this family was present during the early Cretaceous.     

Sex‐biased seed predation in gynodioecious Dianthus superbus var. longicalycinus (Capryophyllaceae) and differential influence of two seed predator species on the floral traits

Gynodioecy, the co‐occurrence of hermaphrodite and female individuals within a species, is maintained by differential reproductive success between sexes. Recently, researchers recognized that not only pollinators but also herbivores are important agents in the evolution and maintenance of gynodioecy, when herbivory is hermaphrodite biased. In this study, we investigated whether there is hermaphrodite‐biased herbivory in a gynodioecious plant, Dianthus superbus var. longicalycinus, and if so, what floral traits influenced hermaphrodite‐biased herbivory. We measured flower morphology (flower diameter, calyx tube length, corolla height and petal width) and phenology of flowers of female individuals, hermaphrodites and gynomonoecious individuals in a natural population. We also investigated seed predation and predator species. At the study site, Sibinia weevils (Curculionidae; Coleoptera) and Coleophora moths (Coleophoridae; Lepidoptera) were common pre‐dispersal seed predators in this species. The weevil appeared early in the flowering season, and weevil predation correlated with flower phenology. Because female individuals did not flower early in the season, weevil predation was less frequent in female individuals. Moth predation correlated with calyx length. The calyx length of flowers of female individuals was smaller than those of hermaphrodites, but a direct comparison of moth predation rates failed to find a significant difference among sex morphs. We found that the two seed predators had different effects on floral traits in D. superbus var. longicalycinus. We suggest that weevil predation contributes to the maintenance of gynodioecy because female individuals successfully escaped weevil predation by flowering late. It remains unclear why flower phenology is different among sex morphs.     

Consequences of flower heliotropism for reproduction in an alpine buttercup (Ranunculus adoneus)

Summary The flowers of the alpine snow buttercup Ranunculus adoneus track the sun's movement from early morning until mid-afternoon. Individual blooms last up to a week: younger female stage flowers show greater solar tracking fidelity than older hermaphrodite or dehisced flowers. Flowers aligned parallel to the sun's rays reach mean internal temperatures several degrees Celsius above ambient air temperature. As a flower's angle of deviation from the sun increases beyond 45 degrees, internal flower temperature is significantly reduced. Fly pollinators are seen disproportionately often on flowers aligned with the sun; this is due, in part, to their greater residence time on tracking flowers. Fly visitation is important to fecundity. When flies were excluded from flowers, some selfed seeds were matured, but total seed production was much less than in either handoutcrossed or open-pollinated flowers. Flowers that were tethered at random angles to prevent solar tracking set fewer, smaller seeds than unmanipulated control flowers. Reductions in seed weight were statistically consistent at different times in the season, but tethering influenced seed number per flower most strongly in the early season. Effects of tethering on maternal reproduction could be due to either pollinator diserimination, post-pollination developmental processes, or both. In a second series of experiments using fluorescent dye particles as pollen analogs, tethered donor flowers dispersed dye to as many recipients as paired control flowers, suggesting that solar tracking fidelity may have little effect upon this component of male reproduction.     

Flower biology of the cactus Coryphantha elephantidens in the tropical dry forest of Central Mexico

Floral traits and sexual systems in angiosperms are strategies that enhance outcrossing within hermaphrodite flowers and among individuals in a population. Sexual systems with unisexual flowers have also evolved among angiosperms, resulting in sex specialization. Furthermore, the interaction of floral traits and floral visitors determines successful plant reproduction. Globose cacti are bee pollinated, and variation in the diversity of their pollinator assemblages is strongly associated with floral phenotype. Our objective was to describe the floral biology of the cactus Coryphantha elephantidens and to determine its relationship with pollinators. Floral traits were studied by direct observations in live and fixed flowers. The breeding system was determined using two estimators based on floral morphology: pollen grains to ovules per flower (P/O) ratio and outcrossing index. Pollination treatments were conducted to determine the mating system. Floral visitors were recorded using direct observation. Flowers of C. elephantidens are variable in color, protandric, herkogamous and nectarless. Estimators of the breeding system indicated xenogamy, which is consistent with the obligate outcrossing revealed by the pollination experiment. Thirty-seven percent of the plants have female flowers that do not produce pollen, making this population functionally gynodioecious. Both fruit and seed set were high compared to other globose cacti. Pollinators included eight species of native bees, a more diverse pollinator assemblage than other globose cacti. Given the high pressure on pollen due to functional gynodioecy, nectarless flowers, an outcrossing mating system, and the necessity of pollinators to set seeds, we concluded that native bees are highly efficient pollinators that play a crucial role in the sexual reproduction of C. elephantidens.     

中国北亚热带油橄榄(城固32)开花生物学特性研究

为研究北亚热带内陆地区油橄榄（Olea europaea L.）开花物候、开花样式及花器官特征,探讨其有性繁殖系统的特点。试验对西秦岭南坡地区油橄榄（城固32）的开花生物学进行研究,统计开花进程,雄花和两性花在花序上的着生位置和数量,测定雄花和两性花形态指标并切片观察,电镜扫描分析雄花与两性花花粉粒。结果表明：（1）在北亚热带北缘内陆气候条件下油橄榄花期集中在5月,单株花期15~20 d,盛花期持续4~6 d;（2）油橄榄具有雄全同株的性系统且花器官较小,雄花及两性花都具有正常发育的雄蕊。通过制作石蜡切片观察,发现两性花的雌蕊正常发育,而雄花只有较小的子房和败育的柱头;（3）雌蕊是两种花型花器官生物量差异的主要部位,两性花雌蕊的生物量明显比雄花大（P<0.01）,树体分化出雄花投入的资源更少;（4）雄花与两性花在花粉量、花粉粒大小上差异不显著,在着生位置上,花序轴顶花全部为两性花,雄花出现在花序轴的中部和基部的几率分别是21.13%和30.77%。油橄榄品种城固32具有典型的雄全同株现象,雄花更倾向于在花序上出现的位置在行使它雄性功能方面并无优势可言,而且雄花花粉粒在数量、活力方面也没有优势,但是雄花的出现增加了花粉数量和P/O值,提高了植株的雄性适合度,雄花的出现也降低了两性花落花导致的树体资源浪费,保障其在资源有限的环境中能够繁殖最大化。     

POLLINATION DYNAMICS IN EPILOBIUM CANUM (ONAGRACEAE): CONSEQUENCES FOR GAMETOPHYTIC SELECTION

Several hand-pollination experiments have shown that pollen tube competition leads to gametophytic selection, and thus affects genotypic and phenotypic characteristics of the next generation. This study is one of the first to quantify natural levels of pollen tube competition. In a population of Epilobium canum, I measured both the amount of pollen deposited on stigmas and the timing of deposition. Approximately 20 tetrads were required for full seed set within fruits. Hummingbirds deposited >20 tetrads at 50–70% of the flowers examined, often in a single load. When pollen arrived in 2 loads, a portion of the ovules within each ovary was probably sired by competing pollen from the second load. Competition may be relatively weak unless at least 30 tetrads compete for ovules. About 20% of the flowers received >30 tetrads in the first load, and 13% acquired > 30 tetrads in 2 loads. The frequency and intensity of pollen tube competition varied among plants. In some styles, 80% of the pollen tubes were excluded from access to ovules, but in others no competition occurred. Further studies of pollination rates and progeny fitness are needed before we can assess the role of pollen tube competition in natural populations. Potential effects of gametophytic selection are discussed.     

Synchronized dichogamy and dioecy in neotropicalLauraceae

In hermaphrodite neotropicalLauraceae a highly evolved dichogamous system is present which represents a kind of temporal dioecy. This system involves the existence of two flower morphs which are characterized by reciprocal phases of receptivity of the stigmas and pollen release. In some genera (Persea, Cinnamomum), nectar is produced as a reward for the flower visitors, while in other genera (Aniba, Clinostemon, Licaria), nectar is absent and pollen seems to be the only reward. This implies that in this case the flowers in the female stage must be deceptive flowers. In dioecious species of the generaOcotea andNectandra, both the male and female flowers attract the visitors with nectar. The pollen-ovule ratio of theLauraceae is comparatively low. — The type of reproductive system that characterizes theLauraceae, comprising functional dioecy, small, inconspicuously coloured flowers, pollination by small bees, and large, one-seeded fruits dispersed by birds, is quite prominent among trees of various families in the tropical lowland forest. The relationship between the different modes of flowering within theLauraceae and the causes for the correlation of their reproductive traits are discussed.     

Sex Expression and the Evolutionary Advantages of Male Flowers in an Andromonoecious Species, Sagittaria guyanensis subsp. lappula (Alismataceae)

Sex expression in Sagittaria guyanensis H.B.K. subsp. lappula (D. Don) Bojin is typically andromonoecious while the other species in the same genus are basically monoecious. The evolutionary advantages of male flowers and hermaphrodite flowers in S. guyanensis subsp. lappula were assessed by measuring sex allocation and pollen movements in two wild populations of the species. Two cultivated populations served as controls. The percentage of male flowers was very low in the two wild populations in Dongxiang, Jiangxi Province and Wuyishan, Fujian Province, viz., 2.48% and 0.96% respectively. In the two cultivated populations, male flower percentage significantly increased when the soil was of higher nutrient content. This indicates that the allocation to male versus female reproduction might change in response to environmental factors. Pollen production per male flower was 4.1 times higher than that of a hermaphrodite flower. The floral shape and size of male and hermaphrodite flowers were similar. No difference was observed between these dimorphological flowers in pollen germination rate in vitro and in the speed of pollen tube growth in vivo. Anthesis was only 4-5 h. Male flowers usually opened 0.5 h earlier than hermaphrodite flowers. An unexpected finding was that no pollen from the male flowers was found on the stigmas of the hermaphrodite flowers, in spite of the occasional visits by insects to both types of flowers in both wild and control populations. A consistent pattern of fruit development was found to exist in open pollinated flowers as well as in flowers that had been bagged. The sex ratios did not have significant influence on fruit set. Approximately 25% of the pistils in a gynoecium failed to develop into fruits because no pollen was deposited on them, indicating that the fruit set of this andromonoecious plant is mainly affected by pollen limitation rather than resource limitation. Reproduction in S. guyanensis subsp. lappula in the habitats was dependent on self-pollination in hermaphrodite flowers. The male flowers in this species might be a potential source of additional pollination and may facilitate cross-pollination. The fact that the flowers of monoecious species in Sagittaria pollinated by a wide diversity of insect visitors may contribute enormously to the diversification of sex expression in this genus.     

Dioecy inHonkenya peploides var.major (Caryophyllaceae)

Sexual difference and pollination system inHonkenya peploides var.major were investigated. The present analyses indicate that sex expression of this plant is functionally dioecious, rather than androdioecious, as described previously. Its flowers are largely pollinated by small bees, hover-flies, flies, and ants. Female flowers were morphologically characterized by conspicuously longer styles and smaller petals, and had no pollen grains, while male flowers had shorter styles, larger petals, and longer filaments, and produced viable pollen grains. Although the male flowers possess many ovules in an ovary, they set fruits at a very low frequency in natural conditions and did not produce fruits by pollination experiments. We suggest that the dioecy of this species has been derived from hermaphrodite through gynodioecy, and not directly from hermaphrodite.     

Status of self-pollen in bee pollination efficiency of white clover (Trifolium repens L.)

Flowers of white clover (Trifolium repens L.) are hermaphrodite and self-incompatible; their cross-pollination depends entirely on insect visitors, mainly bees (Apoidea). Because self-pollination of white clover occurs before flower anthesis, we determined whether selfing affected the pollination efficiency of a honeybee visit. We compared pollen deposition in emasculated and intact flowers following (1) a single honeybee visit, (2) open-pollination for a day and (3) enclosure in a cloth bag to prevent insect visits. In emasculated flowers, open-pollination resulted in more pollen deposited than after one visit (+30%) which is consistent with flowers being visited more than once by pollinators during the course of a day. On intact flowers, saturation of the stigma was achieved after the first visit of a honeybee (near 280 grains) because of self-pollination. Additional visits did not increase pollen deposits, but they improved pollen efficiency in terms of numbers of pollen tubes reaching the ovules. In such a context of easily saturated stigmas, self-pollen does not inhibit cross-pollen activity, but represents a constraint for pollination which demands multiple bee visits to each flower to achieve maximum fertilization. Received: 20 May 1997 / Accepted: 25 October 1997     

Fruit set increases with maleness in the andromonoecious Acacia caven

In long-lived plants, sexual expression can be highly variable in time and space. The extent to which sexual dimorphism of flowers is expressed in the same or in separate individuals, may have implications for the plants’ dependence on pollen vectors, the compatibility of the pollen received, and the potential for setting fruit. Here, we account for variability in sexual expression in the small tree Acacia caven, which produces hermaphrodite and male flowers on the same or in separate individuals. We focus on whether individual sexual expression influences fruit set, which latter has been reported to be extremely low in this species. Individual sexual expression, estimated as the hermaphrodite: male flowers ratio was variable both within and among plants across two reproductive seasons. Though male flower production could be very high (up to 99% of the flowers), all trees produced both flower types within the same individual, thus confirming andromonoecy in this species. More interestingly, hermaphrodite: male flowers ratio had a negative effect on fruit set. This pattern was consistent across two reproductive seasons and suggests that maleness should favor female function. Similar trends found in other plant species with the same sexual system support the hypothesis that male flowers of andromonoecious species may contribute to reduce self-pollen interference compared with hermaphrodite flowers. By favoring the deposition of compatible pollen grains and increasing fruit set, maleness would then help to maintain andromonoecy in this and other andromonoecious species.     

冠果草的性表达状态及其进化含义

通常认为雄花两性花同株是植物性表达两性花进化到雌雄异株的过渡类型之一。慈姑属仅冠果草（Ｓａｇｉｔｔａｒｉａ ｇｕｙａｎｅｎｓｉｓ Ｈ．Ｂ．Ｋ．ｓｕｂｓｐ．ｌａｐｐｕｌａ（Ｄ．Ｄｏｎ）Ｂｏｊｉｎ）的性表达为典型的雄花两性花同株,其他种为雌雄同株。对冠果草野外和实验居群的性配置进行了定量观察。东乡和武夷山居群雄花的比例分别为２．４８％０．９６％。实验表明,栽培条件影响冠果草的性表达：在营养较好的条件下     

The effects of floral display on pollinator visitation vary among populations ofPhacelia linearis (Hydrophyllaceae)

Summary Individual plants in gynodioecious populations ofPhacelia linearis (Hydrophyllaceae) vary in flower gender, flower size, and flower number. This paper reports the effects of variation in floral display on the visitation behaviour of this species' pollinators (mainly pollen-collecting solitary bees) in several natural and three experimental plant populations, and discusses the results in terms of the consequences for plant fitness. The working hypotheses were: (1) that because female plants do not produce pollen, pollen-collecting insects would visit hermaphrodite plants at a higher rate than female plants and would visit more flowers per hermaphrodite than per female; and (2) that pollinator arrival rate would increase with flower size and flower number, the two main components of visual display. These hypotheses were generally supported, but the effects of floral display on pollinator visitation varied substantially among plant populations. Hermaphrodites received significantly higher rates of pollinator arrivals and significantly higher rates of visits to flowers than did females in all experimental populations. Flower size affected arrival rate and flower visit rate positively in natural populations and in two of the three experimental populations. The flower size effect was significant only among female plants in one experimental population, and only among hermaphrodites in another. The effect of flower number on arrival rate was positive and highly significant in natural populations and in all experimental populations. In two out of three experimental populations, insects visited significantly more flowers per hermaphrodite than per female and visited more flowers on many-flowered plants than on few-flowered plants, but neither effect was detected in the third experimental population. Because seed production is not pollen-limited in this species, variation in pollinator visitation behaviour should mainly affect the male reproductive success of hermaphrodite plants. These findings suggest that pollinator-mediated natural selection for floral display inP. linearis varies in space and time.     

Reproductive system and fitness of Vriesea friburgensis,a self‐sterile bromeliad species

Reproductive biology and plant fertility are directly related to many aspects of plant evolution and conservation biology. Vriesea friburgensis is an epiphytic and terrestrial bromeliad endemic to the Brazilian Atlantic rainforest. Hand‐pollination experiments were used to examine the reproductive system in a wild population of V. friburgensis. Plant fertility was assigned considering flower production, fruit and seed set, seed germination, and pollen viability. Self‐sterility observed from spontaneous selfing and manual self‐pollination treatments may be the consequence of late‐acting self‐incompatibility. Hand‐pollination results indicated no pollen limitation in the population studied. Floral biology features such as a few daily open flowers, nectar production, and sugar concentration corroborate hummingbirds as effective pollinators, although bees were also documented as pollinators. Components of fitness such as high flower, fruit, and seed production together with high seed and pollen viability indicate that this wild population is viable. From a conservation point of view, we highlight that this self‐sterile species depends on pollinator services to maintain its population fitness and viability through cross‐pollination. Currently, pollinators are not limited in this population of V. friburgensis. Conversely, the maintenance and continuous conservation of this community is essential for preserving this plant–pollinator mutualism.     

The mechanism of delayed selfing in Collinsia verna (Scrophulariaceae)

Collinsia verna, blue-eyed Mary, has floral attributes of an outcrossing species, yet most flowers readily self-pollinate under greenhouse conditions. Here we describe the mechanism of self-pollination in C. verna via changes in relative positions of the stigma and anthers and late timing of receptivity, resulting in delayed selfing. Each flower contains four anthers that dehisce sequentially over ∼1 wk. Pollen that is not collected by pollinators accumulates in the keel petal and retains high viability (>80% pollen germination) up to the time of corolla abscission. The stigmatic surface does not become receptive until after the third anther dehisces. This overlap in the sexual phases is concurrent with a change in herkogamy during floral development. In most flowers (70%), the stigma has moved to the front of the keel and is positioned near the anthers when the third anther dehisces. Under field conditions, fruiting success of plants within pollinator exclosures was ∼75% of the fruiting success in open-pollinated plants (33% fruiting success via autogamy vs. 44% fruiting success, respectively). Collinsia verna plants in pollinator exclosures exhibit variation in autogamy rates within natural populations (range 0–80%). In addition, only half of naturally pollinated, receptive flowers examined had pollen tubes growing in their styles. In contrast, shortly after corolla abscission, nearly all flowers examined (96%) had pollen tubes in their styles. Thus we find that in C. verna, autogamy occurs late in floral development, which has the potential to provide substantial reproductive assurance, and that individuals vary in their ability to set fruit through this mechanism. We suggest that delayed selfing mechanisms may be overlooked in other species and that variable pollinator availability may play a significant role in the maintenance of mixed mating in species with delayed selfing, such as C. verna.