Call number: LD2668 .T4 1984 L627 / Master of Science

Thesis (doctoral)--Universität Bern, 1904. / Cover title. Includes bibliographical references.

Comprendre la diversité morphologique des fleurs passe par l'étude de son origine moléculaire et développementale et de ses conséquences fonctionnelles et écologiques. Le périanthe est composé d'organes stériles, sépales et pétales, qui jouent un rôle majeur dans le succès reproducteur des plantes pollinisées par les animaux du fait de leur fonction d'attraction.Cette thèse propose une approche multidisciplinaire visant à comprendre l'origine génétique et moléculaire de la diversité morphologique du périanthe et sa signification évolutive, à l'aide du modèle Nigella damascena L. Cette Renonculacée présente un dimorphisme spontané. La forme probablement ancestrale, trouvée en populations naturelles, a un périanthe bipartite composé de cinq sépales pétaloïdes et huit pétales nectarifères. Dans la forme variante, cultivée à des fins d'horticulture, les pétales sont remplacés par un nombre élevé d'organes allant d'une forme proche des sépales à une forme proche des étamines.La première partie de cette thèse est consacrée à l'étude de l'origine développementale, génétique et moléculaire du dimorphisme, par la caractérisation détaillée de la morphologie florale et de son développement dans les deux morphes dans le cadre d'une approche gène candidat. Par analyse d'expression et validation fonctionnelle, nous avons montré que le gène NdAP3-3 est responsable de l'ensemble des aspects du dimorphisme floral de N. damascena, ce qui suggère que ce gène joue un rôle dans l'identité du pétale mais aussi dans l'architecture du méristème, potentiellement via la régulation du nombre d'organes et de la frontière entre périanthe et étamines.La seconde partie de cette thèse concerne l'impact du dimorphisme floral sur le mode de reproduction des deux morphes et leur maintien potentiel. Nous avons caractérisé les stratégies reproductives et la valeur sélective des deux morphes en conditions naturelles dans des populations expérimentales. Le variant sans pétale est peu visité par les pollinisateurs, et se reproduit majoritairement en autogamie. L'analyse de la vigueur de ses descendants suggère une dépression de consanguinité. Par ailleurs, dans notre matériel, il semble que l'allèle donnant le phénotype sans pétale soit lié à un allèle augmentant la valeur sélective. A la lumière de nos résultats, nous discutons les conditions du maintien de ce polymorphisme. / Understanding flower diversity requires on one hand the study of the molecular and developmental origin of floral architecture, and on the other the study of the functional and ecological consequences of flower morphology. A great deal of that diversity can be found at the perianth level which comprises the sepals and petals, sterile and versatile organs that play a major role in the reproductive success of animal pollinated flowering plants through their attractive characteristics.This thesis is the result of a multidisciplinary effort to understand the genetic and molecular origin as well as the evolutionary significance of perianth diversity, using the Nigella damascena L. as a model. This Ranunculaceae species presents a rare naturally occurring floral dimorphism affecting perianth architecture. The putatively ancestral form found in natural populations has a well differentiated bipartite perianth composed of five petaloid sepals and eight nectariferous petals, while the perianth in the alternative apetalous mutant, cultivated for horticultural purpose, has no petals and but is instead composed of numerous organs showing a continuum of forms from outer sepal-like to inner stamen-like.The first part of this thesis was dedicated to the study of the developmental, genetic and molecular origin of this dimorphism, via a detailed characterization of floral morphology and development in both morphs, which laid a foundation for the interpretation of the results of a candidate gene approach. Using expression analysis and functional validation we showed that NdAP3-3 is fully responsible for the complex N. damascena floral dimorphism, suggesting that it plays a role not only in petal identity but also in meristem patterning, possibly through the regulation of perianth organ number and perianth-stamen boundary.The second half of this thesis focused on the impact of the floral dimorphism on the reproduction mode and evolutionary maintenance of the two morphs. We assessed reproduction strategies and reproductive success in the two morphs by studying a polymorphic experimental population in natural conditions. The absence of petals in the mutant form was associated with a qualitative drop in pollinator visitation which resulted in a shift towards selfing. The study of their progeny suggests that selfing had a negative effect on the descendant’s vigor via inbreeding depression. Additionally, in our material, the allele responsible for the apetalous phenotype seems to be linked to a favorable allele increasing fitness. We discuss the mechanisms of the dimorphism maintenance in light of these results.

The genus Clematis L. (Ranunculaceae) was used as a new model group to assess the role of the Himalayan orogeny on generation of biodiversity through investigations of its phylogeny, phylogeography and taxonomy. Although existing checklists include 28 species of Clematis from Nepal, a comprehensive taxonomic revision of available material in herbaria and additional sampling from fieldwork during this study has led to the recognition of 21 species of Clematis in Nepal, including one species (C. kilungensis) not previously recorded from Nepal. Exisiting phylogenetic and taxonomic concepts were tested with the addition of new samples from Nepal. The results highlight the shortcomings of the previous studies which were poorly resolved and indicate the need for a thorough revision of the sectional classification. Despite the increased sampling the results are still equivocal due to poor statistical support along the backbone of the phylogeny. Groups of species in well supported terminal clades are broadly comparable with results from previous studies although there are fewer clearly recognisable and well supported clades. The published dates for the evolution of Clematis were tested and the methodology of the previous study critically reappraised. The results indicate that the genus Clematis is approximately twice as old as previously reported and evolved in the middle Miocene. The phylogeny also demonstrates that, even allowing for poor support for the relationships between groups of species within Clematis, the extant Nepalese species must have multiple independent origins from at least 6 different colonisations. With their occurrence in the Pliocene and Pleistocene, these events are relatively recent in relation to the Himalayan orogeny, and may be linked more to the dispersal ability of Clematis than to the direct effects of the orogeny. Additional Nepalese samples of Koenigia and Meconopsis were added to exisiting datasets and these were reanalysed. The result from Clematis, Koenigia and Meconopsis were appraised in light of the the geocientific literature and previously published phylogeographic studies to create an overview of the drivers behind speciation in the Himalaya.

The relationship between leaf architecture, vein anatomy and phloem ultrastructure, and that of possible routes from mesophyll cells to phloem and potential phloem loading method was investigated using species adapted to the southern African climate. The research was based on the hypothesis of Gamaiei and Van Bel, using nothern hemisphere species only (Gamalei 1985a, b, 1989, 1991, VanBeletal. 1988, Van Bel 1992, 1994, 1996, Van Bel & Gamalei 1991, 1992, Gamalei et al. 1992, 1994, 1996, Van Bel 1992a-c, 1993a, b, 1996). The thesis commenced with a survey ofleafarchitecture of the Ranunculaceae, Apocynaceae and Asclepiadaceae. Anatomical and ultrastructural studies followed. Leaf architecture was described according to Hickey (1973). Within the Ranuncuiaceae, leaf architecture was found to be marginally actinodromous. Venation pattern consisted of a widely spaced reticulum of delicate veins, especially in Ranunculus. Leaf architecture of the Apocynaceae was described as pinnate, camptodromous and brochidodromous. The Asclepiadaceae showed less uniformity in terms of leaf architecture, being pinnate and camptodromous, with mostly brochidodromous and, unexpectedly, eucamptodromous patterns of secondary venation. A predominantly common leaf architecture supported the move to amalgamate the two families. As the less advanced eucamptodromous arrangement could represent a more primitive branch of this huge family, the phylogenetic classification of the new amalgamated family is eagerly awaited for discussion. Allocation of vein order allowed comparisons between species and families to be drawn. Reticulum density and vein order anatomy was used to indicate potential routes from mesophyll to phloem. A definite contrast was obvious between the loose arrangement of mesophyll and veins in the mesic Ranunculus, and the close mesophyll and dense venation of the xeric apocynate and asclepiad species, and was related to habitat. Ultrastructural characteristics of companion cells, together with plasmodesmatal abundance, were considered especially important for the determination of minor vein configuration. Descriptions of plasmodesmatal distribution did not consider functional status. In this thesis, vein structure and ultrastructure were considered in relation to phloem loading, not as a demonstration thereof. All three families were designated minor vein configuration type 2a. Two interesting examples that did not adhere to the familial norm, viz. few plasmodesmata and normal companion cells, occurred in the Asclepiadaceae. Secamone alpinii had abundant aggregated plasmodesmata, forming a potential symplasmic continuum from mesophyll to companion cells. The question of plasmodesmatal functionality remained open. Ceropegia carnosa showed folding of the companion cell membrane, but no accompanying wall ingrowths. The folds were suggested to increase surface area for apoplasmic phloem loading in the noted absence of plasmodesmata. Loading routes and methods suggested were based on anatomical and ultrastructural evidence only. Whilst these results were supported by published data for other species of these families, the prediction of the Gamalei and Van Bel hypothesis did not hold true. The relatively primitive Ranunculaceae were expected to have the least advanced type 1 minor vein configuration, with abundance plasmodesmata providing a symplasmic phloem loading pathway. The relatively advanced Apocynaceae and Asc1epiadaceae were predicted to have the most progressive minor vein configuration, type 2b, with specialised transfer cells to maximise apoplasrnic uptake. As families with type 2a minor vein configurations, the Ranunculaceae were more advanced than expected and the Apocynaceae and Asc1epiadaceae less so.

京都大学 / 0048 / 新制・課程博士 / 博士(人間・環境学) / 甲第21875号 / 人博第904号 / 新制||人||215(附属図書館) / 2018||人博||904(吉田南総合図書館) / 京都大学大学院人間・環境学研究科相関環境学専攻 / (主査)教授 瀬戸口 浩彰, 教授 加藤 眞, 教授 市岡 孝朗, 准教授 西川 完途 / 学位規則第4条第1項該当 / Doctor of Human and Environmental Studies / Kyoto University / DGAM

Comprendre la diversité morphologique des fleurs passe par l'étude de son origine moléculaire et développementale et de ses conséquences fonctionnelles et écologiques. Le périanthe est composé d'organes stériles, sépales et pétales, qui jouent un rôle majeur dans le succès reproducteur des plantes pollinisées par les animaux du fait de leur fonction d'attraction.Cette thèse propose une approche multidisciplinaire visant à comprendre l'origine génétique et moléculaire de la diversité morphologique du périanthe et sa signification évolutive, à l'aide du modèle Nigella damascena L. Cette Renonculacée présente un dimorphisme spontané. La forme probablement ancestrale, trouvée en populations naturelles, a un périanthe bipartite composé de cinq sépales pétaloïdes et huit pétales nectarifères. Dans la forme variante, cultivée à des fins d'horticulture, les pétales sont remplacés par un nombre élevé d'organes allant d'une forme proche des sépales à une forme proche des étamines.La première partie de cette thèse est consacrée à l'étude de l'origine développementale, génétique et moléculaire du dimorphisme, par la caractérisation détaillée de la morphologie florale et de son développement dans les deux morphes dans le cadre d'une approche gène candidat. Par analyse d'expression et validation fonctionnelle, nous avons montré que le gène NdAP3-3 est responsable de l'ensemble des aspects du dimorphisme floral de N. damascena, ce qui suggère que ce gène joue un rôle dans l'identité du pétale mais aussi dans l'architecture du méristème, potentiellement via la régulation du nombre d'organes et de la frontière entre périanthe et étamines.La seconde partie de cette thèse concerne l'impact du dimorphisme floral sur le mode de reproduction des deux morphes et leur maintien potentiel. Nous avons caractérisé les stratégies reproductives et la valeur sélective des deux morphes en conditions naturelles dans des populations expérimentales. Le variant sans pétale est peu visité par les pollinisateurs, et se reproduit majoritairement en autogamie. L'analyse de la vigueur de ses descendants suggère une dépression de consanguinité. Par ailleurs, dans notre matériel, il semble que l'allèle donnant le phénotype sans pétale soit lié à un allèle augmentant la valeur sélective. A la lumière de nos résultats, nous discutons les conditions du maintien de ce polymorphisme.