Ph.D. / Cussonia Thunb. is a genus of 21 species (including one that is currently undescribed) found in grasslands, woodlands and forests of sub- Saharan Africa, the Arabian Peninsula (Yemen) and the Comoro Islands. The genus generally has soft brittle branches, a fleshy underground tuber and fleshy roots. Despite these features, Cussonia is not found in the dry and arid regions of Africa. They are generally trees or shrubs, and less often lianes (C. thyrsiflora Thunb.) or suffrutices (C. corbisieri De Wild.). The tree forms usually grow between 4 to 20 m in height; however C. zimmermannii Harms can grow up to 45 m tall. Cussonia corbisieri has a large woody underground caudex with multiple 1 m tall, herbaceous stems. Probably the most distinguishing feature of this genus is the shape and size of the leaves. The palmately compound leaves are carried (in most cases) on the ends of the branches. The simple palmatisect leaves, such as in C. arborea Hochst. ex A.Rich. or C. natalensis Sond., are not subdivided into smaller leaflets, while the compound leaves increase in complexity from mono-palmately compound to multi-palmately compound. The multi-palmate compound leaves are carried on long petioles (up to 700 mm) having up to nine leaflets (up to 500 mm long), with each leaflet having two or more articulations bearing secondary leaflets (up to 400 mm long). These secondary leaflets can be subdivided into tertiary leaflets (up to 200 mm long).

Thesis (Ph.D)--Virginia Commonwealth University, 2009. / Prepared for: Dept. of Biology. Title from resource description page. Includes bibliographical references.

Obligate shade plants possess adaptations that enable them to photosynthesise in the low light environment of the forest floor. Adaptations that facilitate light scavenging may compromise capacity for high rates of photosynthesis. This study compares the responses of obligate shade and facultative shade plant species upon exposure to elevated light. The obligate shade plants were two commercially grown medicinal herb species of ginseng, Panax ginseng C.A. Meyer and Panax quinquefolius L.; and goldenseal - Hydrastis canadensis L. Comparison was made to Arabidopsis thaliana and Pisum sativum L. as facultative shade species. Panax ginseng (Korean ginseng) and Panax quinquefolius (American ginseng) are obligate shade plants found in broadleaf forests of Eastern Asia and North America, respectively. Studies on these plants have shown optimal growth at light intensities between 200-300 [mu]mol photons. m⁻�. s⁻�, or 10-15% of full sunlight, and at intensities greater than 500 [mu]mol photons. m⁻�. s⁻� characteristic photoinbibitory symptoms develop. An atypical response to methyl viologen in photosynthetic electron transport assays was observed in ginseng in both isolated thylakoid membranes and whole leaves. No correlation was found between detectable superoxide dismutase activity and altered methyl viologen reactions. In a mutagenesis study using the model cyanobacterium Synechocystis sp. PCC 6803, a unique amino acid residue in the terminal electron acceptor PsaC, found only in ginseng, was changed and found to have no effect on methyl viologen reactions. Electron transfer to methyl viologen was examined in both isolated thylakoid membranes and whole leaves using chlorophyll a fluorescence and the apparent ability for methyl viologen to act as an electron acceptor was observed to differ between ginseng species. Obligate shade species were observed to possess alternate pools of photosystem II centres that potentially provide a mechanism to maximise photosynthetic gain under low light and during short periods of increased illumination. In experiments designed to identify physiological processes that contribute to increased susceptibility to photoinhibition in obligate shade plants, responses were observed and characterised following a moderate increase in illumination (140 to 400 [mu]mol photons. m⁻� . s⁻�) using chlorophyll a fluorescence induction curve analysis. The obligate shade species exhibited varied responses to elevated light and showed increased susceptibility, to photoinhibition. Photoprotective non-photochemical dissipative capacity was quantified and found to be comparable between all species studied.

Thesis (M.S.)--University of Missouri-Columbia, 2006. / Includes bibliographical references (p. 157-167). Also available in PDF format via the Internet.

Obligate shade plants possess adaptations that enable them to photosynthesise in the low light environment of the forest floor. Adaptations that facilitate light scavenging may compromise capacity for high rates of photosynthesis. This study compares the responses of obligate shade and facultative shade plant species upon exposure to elevated light. The obligate shade plants were two commercially grown medicinal herb species of ginseng, Panax ginseng C.A. Meyer and Panax quinquefolius L.; and goldenseal - Hydrastis canadensis L. Comparison was made to Arabidopsis thaliana and Pisum sativum L. as facultative shade species. Panax ginseng (Korean ginseng) and Panax quinquefolius (American ginseng) are obligate shade plants found in broadleaf forests of Eastern Asia and North America, respectively. Studies on these plants have shown optimal growth at light intensities between 200-300 [mu]mol photons. m⁻�. s⁻�, or 10-15% of full sunlight, and at intensities greater than 500 [mu]mol photons. m⁻�. s⁻� characteristic photoinbibitory symptoms develop. An atypical response to methyl viologen in photosynthetic electron transport assays was observed in ginseng in both isolated thylakoid membranes and whole leaves. No correlation was found between detectable superoxide dismutase activity and altered methyl viologen reactions. In a mutagenesis study using the model cyanobacterium Synechocystis sp. PCC 6803, a unique amino acid residue in the terminal electron acceptor PsaC, found only in ginseng, was changed and found to have no effect on methyl viologen reactions. Electron transfer to methyl viologen was examined in both isolated thylakoid membranes and whole leaves using chlorophyll a fluorescence and the apparent ability for methyl viologen to act as an electron acceptor was observed to differ between ginseng species. Obligate shade species were observed to possess alternate pools of photosystem II centres that potentially provide a mechanism to maximise photosynthetic gain under low light and during short periods of increased illumination. In experiments designed to identify physiological processes that contribute to increased susceptibility to photoinhibition in obligate shade plants, responses were observed and characterised following a moderate increase in illumination (140 to 400 [mu]mol photons. m⁻� . s⁻�) using chlorophyll a fluorescence induction curve analysis. The obligate shade species exhibited varied responses to elevated light and showed increased susceptibility, to photoinhibition. Photoprotective non-photochemical dissipative capacity was quantified and found to be comparable between all species studied.

The genus Schefflera is the largest in Araliaceae, with approximately 900 species. Recent studies have shown that Schefflera is polyphyletic and represents no fewer than five distinct clades, each corresponding to a specific geographic region including Asia, continental Africa and Madagascar, Melanesia, the Neotropics, and a small clade distributed throughout several islands in the insular Pacific Ocean. The Afro-Malagasy clade contains nearly 50 species distributed throughout tropical, sub-Saharan Africa, Madagascar, the Comoros, and the Seychelles islands. Previous studies have suggested that this group is monophyletic, identifying two smaller subclades within Afro-Malagasy Schefflera corresponding roughly to informal groups identified as “Meiopanax” and “Sciodaphyllum” on the basis of morphology. Using sequence data from nuclear rDNA spacers and plastid markers derived from 32 of the 48 currently circumscribed species of Afro-Malagasy Schefflera, this study tested the monophyly of Afro-Malagasy Schefflera and of each of its two proposed subclades. Trees based on this molecular data were used to examine patterns of morphological evolution and biogeography among species in the clade. Results support the monophyly of Afro-Malagasy Schefflera and both subclades, which correspond closely to “Meiopanax” and “Sciodaphyllum” which are herein referred to as Neocussonia and Astropanax, respectively. Additional interspecific relationships were examined, which provides evidence for hybridization among several species. Schefflera myriantha, the most widely distributed species of Afro-Malagasy Schefflera, is paraphyletic with respect to two other species, S. humblotiana and S. monophylla. Many morphological features historically used to distinguish species of Afro-Malagasy Schefflera appear to be evolutionarily labile, with a history of gains and losses (e.g., reduction in leaflet number, which occurs independently in both subclades). Biogeographic analyses suggest an African ancestry for the entire Afro-Malagasy Schefflera clade, and for both subclades, with two independent divergence events to Madagascar.

Schefflera is the largest genus in the angiosperm family Araliaceae, with about 900 species, of which c. 300 belong to five subgeneric groups in the Neotropical region. Previous phylogenetic studies of Schefflera have been limited to a small number of species from this region, and very little is know about phylogenetic relationships in the Brazilian-centered Didymopanax group of this genus. Therefore, to gain a better understanding of the diversity and evolution of the Didymopanax group of Schefflera, I investigated the systematics of these plants in the broader context of the entire Neotropical clade. The main goals were (1) to investigate pollen diversity in Neotropical species of Schefflera; (2) to test the monophyly of these species; (3) to provide a taxonomic revision for species of the Didymopanax group of Schefflera; and (4) to investigate evolutionary relationships within the Didymopanax group. Pollen morphology exhibits an uneven variability across Neotropical Schefflera. For example, pollen characters support the distinctiveness of the Didymopanax group from all remaining groups. Moreover, S. tremula has a distinctive pollen morphology compared to remaining species of the Sciodaphyllum group. However, among the remaining groups of Neotropical Schefflera, pollen characters are less distinctive. Molecular phylogenetic analyses have confirmed the monophyly of the Neotropical species of Schefflera, and helped to identify four major clades. One of these clades includes subclades representing the Didymopanax and Crepinella groups, while another clade includes all species from groups Cotylanthes and Sciodaphyllum, excluding Schefflera tremula, a finding that corroborates pollen data. In the formal taxonomic revision of Didymopanax, 37 species are recognized, together with three insufficiently known species. The revision also includes updated species circumscriptions and nomenclatural adjustments for 26 names. Phylogenetic analyses among Didymopanax species recovered four morphologically and geographically coherent clades (Atlantic Forest, Imeri, Five-carpellate and Savannic clades), but their phylogenetic inter-relationships were generally weakly supported. Poorly resolved relationships in the Savannic clade suggests a rapid diversification in the campos rupestres vegetation, which accounts for the greatest species richness in the group. The presence of multiple Didymopanax lineages in the Amazonian and Atlantic forests corroborates that these regions may be composite biogeographic areas.

Thesis (doctoral)--Universität Bern, 1913. / Includes bibliographical references.

L’intérêt des substances d’origine naturelle, potentiellement anti-tumorales nous a amené à nous intéresser aux saponines triterpéniques et stéroïdiques de plantes issues de la biodiversité africaine de la famille des Araliaceae et des Dracaenaceae. En effet, des études antérieures menées sur quelques plantes de ces deux familles ont conduit à l’obtention de molécules complexes et originales possédant d’excellentes propriétés cytotoxiques, immuno-modulatrices, anti-inflammatoires. Au vu de ces résultats nous avons entrepris des investigations pharmaco-chimiques sur Cussonia arborea (Araliaceae), Dracaena deisteliana et Dracaena arborea (Dracaenaceae), plantes médicinales couramment utilisées en pharmacopée traditionnelle africaine pour traiter différentes maladies. Les travaux menés ont conduit à l’isolement de 31 composés purs en utilisant les différentes techniques analytiques du laboratoire notamment les diverses techniques de chromatographie liquide successive à pression atmosphérique, moyenne pression et flash chromatographie sur silice en phase normale et en phase inverse. Les structures ont été déterminées par les méthodes de spectrométrie de masse en source FAB et de spectroscopie de RMN 1D et 2D (COSY, TOCSY, NOESY, HMBC et HSQC). Parmi les 07 composés purs obtenus des écorces de Cussonia arborea, 5 sont des nouvelles saponines triterpéniques dont un dérivé de l’acide ursolique, un dérivé de l’hédéragénine et trois dérivés de l’acide oléanolique, tous disubstitués en position 3 et 28 par des chaînes oligosaccharidiques. 13 composés purs sont obtenus à partir des feuilles de Cussonia arborea, dont 7 nouvelles saponines triterpéniques dérivés de l’acide ursolique, de l’acide 23-hydroxyursolique, de l’hédéragénine et de l’acide oléanolique dont 04 d’entre elles sont obtenues sous forme de mélanges inséparables d’isomères acide oléanolique/acide ursolique et hédéragénine/acide 23-hydroxyursolique. A partir des écorces de Dracaena arborea et des tiges de Dracaena deisteliana, nous avons isolé et caractérisé 10 saponines stéroïdiques dont 4 nouvelles et une sapogénine. Les activités de certains de ces produits purs ont été évaluées sur deux lignées de cellules cancéreuses coliques humaines HCT 116 et HT-29. / The interest of the substances from natural origin, potentially antitumor led us to interest in triterpenoid and steroidal saponins of plants from the African biodiversity belonging to the Araliaceae and Dracaenaceae families of plants. Indeed, of the former studies undertaken on some plants of these two families led to obtaining complex and original molecules having excellent cytotoxic, immuno-modulating, anti-inflammatory properties. Within sight of these results we undertook pharmaco-chemical investigations on Cussonia arborea (Araliaceae), Dracaena deisteliana, and Dracaena arborea (Dracaenaceae), medicinal plants usually used in african traditional pharmacopeia to treat various diseases. The work led to the isolation of 31 pure compounds by using the various analytical techniques in particular the various chromatography techniques (CC, MPLC, TLC, flash) on silica gel, normal and reversed phases. The structures were determined by the methods of mass spectrometry (FAB, ESI, IE) and 1D (1H and 13C) and 2D (COSY, TOCSY, NOESY, HMBC and HSQC) NMR spectroscopy. Among the 07 pure compounds obtained of the barks of Cussonia arborea, 5 are new triterpenoid saponins derivatives of ursolic acid, hederagenin and three derived from the acid oleanolic, all disubstituted in position 3 and 28 by oligosaccharidic chains. 13 pure compounds were obtained from leaves of Cussonia arborea, seven of which are new triterpenoid saponins derivatives of oleanolic acid, ursolic acid, hederagenin and 23-hydroxyursolic acid of which four were obtained as mixtures of isomers oleanolic acid/ursolic acid and hederagenin/23-hydroxyursolic acid. From the bark of Dracaena arborea and stem of Dracaena deisteliana, we isolated and characterized ten steroidal saponins including 4 new and sapogenin. The activities of some of these pure products were evaluated on two cancerous lines human colic cells HCT 116 and HT-29.

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.