Academic literature on the topic 'Dichogamy'

Duodichogamy and multi-cycle dichogamy are rare forms of temporal separation of staminate and pistillate reproductive functions in angiosperms. We studied the floral phenology, breeding system and pollination of Bridelia retusa (L.) A.Juss. and Bridelia moonii Thwaites, with a particular focus on the alternation of sexual phases to determine variation in their dichogamy. Three dichogamy patterns were identified in B. retusa, including one-cycle dichogamy (staminate → pistillate or pistillate → staminate), duodichogamy (staminate → pistillate → staminate), multi-cycle dichogamy (repeated flowering cycles alternating between staminate and pistillate) and pure staminates. Except for multi-cycle dichogamy, the other dichogamy patterns were prevalent in B. moonii. This study speculated floral mimicry system in Bridelia associated with blowflies.

We developed a novel scoring system to assess spring phenology in a northern red oak (Quercus rubra L.) clonal seed orchard. The system was used to score from 304 to 364 ramets for three reproductive seasons and to place clones into early, intermediate, and late phenology classes. Although the absolute number of clones in each phenological class changed from year to year, the overall order of clonal flowering was highly stable (rs = 0.67, p < 0.001). Early-flowering clones flowered significantly longer than later flowering clones in all 3 years. Dichogamy was present in the orchard, with male flowers of a clone emerging from 1.4 to 3.0 d sooner than its female flowers. Mean dichogamy values for individual clones ranged from 0.0 to 4.9 (± 1.3) d. Year strongly influenced a clone’s dichogamy value (F = 6.0, p = 0.004), whereas genotype had no influence. The mean overall phenological synchronicity for the 3 years of observations was 0.30 ± 0.01 or about 30% overlap between the time when females were receptive and males were shedding pollen. This study represents the first effort to quantify phenology in an artificial population of northern red oak, and it provides a snapshot of the current relationship between temperature, phenology, and floral synchronization.

Abstract The cinnamon flowers exhibit protogynous dichogamy with 2 flower types “Type A” and “Type B” which first flowers during morning and evening respectively. This floral cycle causes a temporal barrier to the maintenance of elite breeding material and for hybridization with desired parents. Determination of variation in flower and inflorescence morphology can shed light on functional diversity in “Type A” and “Type B” flowers. In order to study these variations, a survey of cultivated cinnamon lands and wild habitats was conducted in fifteen locations in the Matara district. Peduncle length (PDL), flower length (FL), flower width (FW), and floral tube length (FTL) varied among cinnamon accessions collected. The variation in tepal shape was distinct: the two whorls of tepals of a single flower exhibited two shapes. The current investigation of differences in inflorescence, floral morphology and floral abnormalities of Cinnamomum verum provides information about their diversity, and recommends molecular analysis to further determine the genetic basis of two flower types in progynous dichogamy.

Although dichogamy is a prevailing feature of the angiosperms, the simultaneous change from male to female phases among hermaphrodite flowers within a plant (i.e., synchronous protandry) has been reported for only a few families (e.g., Araliaceae, Umbelliferae). Here we present an example of synchronous protandry at the ramet level in the Alstroemeriaceae. Dichogamy was analyzed in clonal Alstroemeria aurea at the flower, ramet, and at the whole flowering patch level. Alstroemeria aurea is self-compatible but totally dependent on biotic agents for pollen transfer. There was evidence of strong inbreeding depression expressed during seed development. Comparisons of seed set in open-pollinated flowers with those obtained after hand selfing and outcrossing resulted in a selfing rate of 0.3. At the flower level protandry was complete. The male phase lasted about 4 days and the female phase lasted about 3 days. Between the female and male phase, there was an approximately 1-day long "neuter" phase. Flowering ramets produce a terminal inflorescence bearing one or more whorls of flowers. Within a ramet, flowers of the same order opened within a period of 1–2 days, and male and female phases of different flowers did not overlap. When inflorescences held two whorls of flowers, the ramet went through two alternating non-overlapping male–female cycles. Using spatial autocorrelation techniques, we found little evidence for pairs of neighboring ramets expressing the same sexual phase beyond random expectations at any scale ranging between 0.25 to 15 m. By ensuring pollen interchange between flowering ramets, synchronized protandry at the ramet level could be an important feature in reducing selfing in A. aurea. Key words: Alstroemeria aurea, dichogamy, synchronous protandry, inbreeding depression.

Thirteen cultivars of pecan [Carya illinoinensis (Wangenh.) K. Koch] were monitored for bud break, pollen shed and stigma receptivity for 4 years at LSU Pecan Station, Robson, LA. Cultivars were generally consistent in displaying clear patterns of protogyny or protandry, although patterns were uncertain for some cultivars in some years. Mean dates of cultivar phenology varied significantly by year. Years with warm winter and spring temperatures had earlier seasons of growth and flowering than years with cooler temperatures. The duration of pollen shed and stigma receptivity varied between years. Protogynous cultivars, as a group, had greater bloom overlap than protandrous cultivars, although overlap varied between years for both dichogamy classes. The sequence of cultivar flowering relative to other cultivars varied between years, resulting in variable amounts of bloom overlap between cultivars in different years.

A renewed focus on generalised pollinator systems has inspired a conceptual framework which highlights that spatial and temporal interactions among plants and their assemblage of pollinators can vary across the individual, population, regional and species levels. Pollination is clearly a dynamic interaction, varying in the number and interdependence of participants and the strength of the outcome of the interaction. Therefore, the role of variation in pollination is fundamental for understanding ecological dynamics of plant populations and is a major factor in the evolution and maintenance of generalised and specialised pollination systems. My study centred on these basic concepts by addressing the following questions: (1) How variable are pollinators in a generalised pollination system? To what degree do insect visitation rates and assemblage composition vary spatially among populations and temporally among flowering seasons? (2) How does variation in pollinators affect plant reproductive success? I chose to do this using a model system, Trachymene incisa subsp. incisa (Apiaceae), which is a widespread Australian herbaceous species with simple white flowers grouped into umbels that attract a high diversity of insect visitors. The Apiaceae are considered to be highly generalist in terms of pollination, due to their simple and uniform floral display and easily accessible floral rewards. Three populations of T. incisa located between 70 km and 210 km apart were studied over 2-3 years. The few studies investigating spatial and temporal variation simultaneously over geographic and yearly/seasonal scales indicate that there is a trend for more spatial than temporal variation in pollinators of generalist-pollinated plants. My study showed both spatial and temporal variation in assemblage composition among all populations and variation in insect visitation rates, in the form of a significant population by year interaction. However, removing ants from the analyses to restrict the assemblage to flying insects and the most likely pollinators, resulted in a significant difference in overall visitation rate between years but no difference in assemblage composition between the Myall Lakes and Tomago populations. These results indicate more temporal than spatial variation in the flying insect visitor assemblage of T. incisa. Foraging behaviour provides another source of variation in plant-pollinator interactions. Trachymene incisa exhibits umbels that function as either male or female at any one time and offer different floral rewards in each phase. For successful pollination, pollinators must visit both male and female umbels during a foraging trip. Insects showed both preferences and non-preferences for umbel phases in natural patches where the gender ratio was male biased. In contrast, insects showed no bias in visitation during a foraging trip or in time spent foraging on male and female umbels in experimental arrays where the gender ratio was equal. Pollinator assemblages consisting of a mixture of different pollinator types coupled with temporal variation in the assemblages of populations among years maintains generalisation at the population/local level. In addition, spatial variation in assemblages among populations maintains generalisation at the species level. Fire alters pollination in T. incisa by shifting the flowering season and reducing the abundance of flying insects. Therefore, fire plays an important role in maintaining spatial and temporal variation in this fire-prone system. Although insect pollinators are important in determining the mating opportunities of 90% of flowering plant species worldwide, few studies have looked at the effects of variation in pollinator assemblages on plant reproductive success and mating. In T. incisa, high insect visitation rates do not guarantee high plant reproductive success, indicating that the quality of visit is more important than the rate of visitation. This is shown by comparing the Agnes Banks and Myall Lakes populations in 2003: Agnes Banks received the highest visitation rate from an assemblage dominated by ants but produced the lowest reproductive output, and Myall Lakes received the lowest visitation rate by an assemblage dominated by a native bee and produced the highest seedling emergence. Interestingly, populations with different assemblage composition can produce similar percentage seed set per umbel. However, similar percentage seed set did not result in similar percentage seedling emergence. Differences among years in reproductive output (total seed production) were due to differences in umbel production (reproductive effort) and proportion of umbels with seeds, and not seed set per umbel. Trachymene incisa is self-compatible and suffers weak to intermediate levels of inbreeding depression through early stages of the life cycle when seeds are self-pollinated and biparentally inbred. Floral phenology, in the form of synchronous protandry, plays an important role in avoiding self-pollination within umbels and reducing the chance of geitonogamous pollination between umbels on the same plant. Although pollinators can increase the rate of inbreeding in T. incisa by foraging on both male and female phase umbels on the same plant or closely related plants, most consecutive insect movements were between plants not located adjacent to each other. This indicates that inbreeding is mostly avoided and that T. incisa is a predominantly outcrossing species, although further genetic analyses are required to confirm this hypothesis. A new conceptual understanding has emerged from the key empirical results in the study of this model generalised pollination system. The large differences among populations and between years indicate that populations are not equally serviced by pollinators and are not equally generalist. Insect visitation rates varied significantly throughout the day, highlighting that sampling of pollinators at one time will result in an inaccurate estimate and usually underestimate the degree of generalisation. The visitor assemblage is not equivalent to the pollinator assemblage, although non-pollinating floral visitors are likely to influence the overall effectiveness of the pollinator assemblage. Given the high degree of variation in both the number of pollinator species and number of pollinator types, I have constructed a model which includes the degree of ecological and functional specialisation of a plant species on pollinators and the variation encountered across different levels of plant organisation. This model describes the ecological or current state of plant species and their pollinators, as well as presenting the patterns of generalisation across a range of populations, which is critical for understanding the evolution and maintenance of the system. In-depth examination of pollination systems is required in order to understand the range of strategies utilised by plants and their pollinators, and I advocate a complete floral visitor assemblage approach to future studies in pollination ecology. In particular, future studies should focus on the role of introduced pollinators in altering generalised plant-pollinator systems and the contribution of non-pollinating floral visitors to pollinator assemblage effectiveness. Comparative studies involving plants with highly conserved floral displays, such as those in the genus Trachymene and in the Apiaceae, will be useful for investigating the dynamics of generalised pollination systems across a range of widespread and restricted species.

Doctor of Philosophy (PhD)
A renewed focus on generalised pollinator systems has inspired a conceptual framework which highlights that spatial and temporal interactions among plants and their assemblage of pollinators can vary across the individual, population, regional and species levels. Pollination is clearly a dynamic interaction, varying in the number and interdependence of participants and the strength of the outcome of the interaction. Therefore, the role of variation in pollination is fundamental for understanding ecological dynamics of plant populations and is a major factor in the evolution and maintenance of generalised and specialised pollination systems. My study centred on these basic concepts by addressing the following questions: (1) How variable are pollinators in a generalised pollination system? To what degree do insect visitation rates and assemblage composition vary spatially among populations and temporally among flowering seasons? (2) How does variation in pollinators affect plant reproductive success? I chose to do this using a model system, Trachymene incisa subsp. incisa (Apiaceae), which is a widespread Australian herbaceous species with simple white flowers grouped into umbels that attract a high diversity of insect visitors. The Apiaceae are considered to be highly generalist in terms of pollination, due to their simple and uniform floral display and easily accessible floral rewards. Three populations of T. incisa located between 70 km and 210 km apart were studied over 2-3 years. The few studies investigating spatial and temporal variation simultaneously over geographic and yearly/seasonal scales indicate that there is a trend for more spatial than temporal variation in pollinators of generalist-pollinated plants. My study showed both spatial and temporal variation in assemblage composition among all populations and variation in insect visitation rates, in the form of a significant population by year interaction. However, removing ants from the analyses to restrict the assemblage to flying insects and the most likely pollinators, resulted in a significant difference in overall visitation rate between years but no difference in assemblage composition between the Myall Lakes and Tomago populations. These results indicate more temporal than spatial variation in the flying insect visitor assemblage of T. incisa. Foraging behaviour provides another source of variation in plant-pollinator interactions. Trachymene incisa exhibits umbels that function as either male or female at any one time and offer different floral rewards in each phase. For successful pollination, pollinators must visit both male and female umbels during a foraging trip. Insects showed both preferences and non-preferences for umbel phases in natural patches where the gender ratio was male biased. In contrast, insects showed no bias in visitation during a foraging trip or in time spent foraging on male and female umbels in experimental arrays where the gender ratio was equal. Pollinator assemblages consisting of a mixture of different pollinator types coupled with temporal variation in the assemblages of populations among years maintains generalisation at the population/local level. In addition, spatial variation in assemblages among populations maintains generalisation at the species level. Fire alters pollination in T. incisa by shifting the flowering season and reducing the abundance of flying insects. Therefore, fire plays an important role in maintaining spatial and temporal variation in this fire-prone system. Although insect pollinators are important in determining the mating opportunities of 90% of flowering plant species worldwide, few studies have looked at the effects of variation in pollinator assemblages on plant reproductive success and mating. In T. incisa, high insect visitation rates do not guarantee high plant reproductive success, indicating that the quality of visit is more important than the rate of visitation. This is shown by comparing the Agnes Banks and Myall Lakes populations in 2003: Agnes Banks received the highest visitation rate from an assemblage dominated by ants but produced the lowest reproductive output, and Myall Lakes received the lowest visitation rate by an assemblage dominated by a native bee and produced the highest seedling emergence. Interestingly, populations with different assemblage composition can produce similar percentage seed set per umbel. However, similar percentage seed set did not result in similar percentage seedling emergence. Differences among years in reproductive output (total seed production) were due to differences in umbel production (reproductive effort) and proportion of umbels with seeds, and not seed set per umbel. Trachymene incisa is self-compatible and suffers weak to intermediate levels of inbreeding depression through early stages of the life cycle when seeds are self-pollinated and biparentally inbred. Floral phenology, in the form of synchronous protandry, plays an important role in avoiding self-pollination within umbels and reducing the chance of geitonogamous pollination between umbels on the same plant. Although pollinators can increase the rate of inbreeding in T. incisa by foraging on both male and female phase umbels on the same plant or closely related plants, most consecutive insect movements were between plants not located adjacent to each other. This indicates that inbreeding is mostly avoided and that T. incisa is a predominantly outcrossing species, although further genetic analyses are required to confirm this hypothesis. A new conceptual understanding has emerged from the key empirical results in the study of this model generalised pollination system. The large differences among populations and between years indicate that populations are not equally serviced by pollinators and are not equally generalist. Insect visitation rates varied significantly throughout the day, highlighting that sampling of pollinators at one time will result in an inaccurate estimate and usually underestimate the degree of generalisation. The visitor assemblage is not equivalent to the pollinator assemblage, although non-pollinating floral visitors are likely to influence the overall effectiveness of the pollinator assemblage. Given the high degree of variation in both the number of pollinator species and number of pollinator types, I have constructed a model which includes the degree of ecological and functional specialisation of a plant species on pollinators and the variation encountered across different levels of plant organisation. This model describes the ecological or current state of plant species and their pollinators, as well as presenting the patterns of generalisation across a range of populations, which is critical for understanding the evolution and maintenance of the system. In-depth examination of pollination systems is required in order to understand the range of strategies utilised by plants and their pollinators, and I advocate a complete floral visitor assemblage approach to future studies in pollination ecology. In particular, future studies should focus on the role of introduced pollinators in altering generalised plant-pollinator systems and the contribution of non-pollinating floral visitors to pollinator assemblage effectiveness. Comparative studies involving plants with highly conserved floral displays, such as those in the genus Trachymene and in the Apiaceae, will be useful for investigating the dynamics of generalised pollination systems across a range of widespread and restricted species.