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Flowers on the tree of life
\"Genetic and molecular studies have recently come to dominate botanical research at the expense of more traditional morphological approaches. This broad introduction to modern flower systematics demonstrates the great potential that floral morphology has to complement molecular data in phylogenetic and evolutionary investigations. Contributions from experts in floral morphology and evolution take the reader through examples of how flowers have diversified in a large variety of lineages of extant and fossil flowering plants. They explore angiosperm origins and the early evolution of flowers and analyse the significance of morphological characters for phylogenetic reconstructions on the tree of life. The importance of integrating morphology into modern botanical research is highlighted through case studies exploring specific plant groups where morphological investigations are having a major impact. Examples include the clarification of phylogenetic relationships and understanding the significance and evolution of specific floral characters, such as pollination mechanisms and stamen and carpel numbers\"-- Provided by publisher.
Book
Evolving Ideas on the Origin and Evolution of Flowers: New Perspectives in the Genomic Era
The origin of the flower was a key innovation in the history of complex organisms, dramatically altering Earth’s biota. Advances in phylogenetics, developmental genetics, and genomics during the past 25 years have substantially advanced our understanding of the evolution of flowers, yet crucial aspects of floral evolution remain, such as the series of genetic and morphological changes that gave rise to the first flowers; the factors enabling the origin of the pentamerous eudicot flower, which characterizes ∼70% of all extant angiosperm species; and the role of gene and genome duplications in facilitating floral innovations. A key early concept was the ABC model of floral organ specification, developed by Elliott Meyerowitz and Enrico Coen and based on two model systems, Arabidopsis thaliana and Antirrhinum majus. Yet it is now clear that these model systems are highly derived species, whose molecular genetic-developmental organization must be very different from that of ancestral, as well as early, angiosperms. In this article, we will discuss how new research approaches are illuminating the early events in floral evolution and the prospects for further progress. In particular, advancing the next generation of research in floral evolution will require the development of one or more functional model systems from among the basal angiosperms and basal eudicots. More broadly, we urge the development of “model clades” for genomic and evolutionary-developmental analyses, instead of the primary use of single “model organisms.” We predict that new evolutionary models will soon emerge as genetic/genomic models, providing unprecedented new insights into floral evolution.
Journal Article
Development and evolution of extreme synorganization in angiosperm flowers and diversity
Apocynaceae and Orchidaceae are two angiosperm families with extreme flower synorganization. They are unrelated, the former in eudicots, the latter in monocots, but they converge in the formation of pollinia and pollinaria, which do not occur in any other angiosperm family, and for which extreme synorganization of floral organs is a precondition. In each family extensive studies on flower development and evolution have been performed; however, newer comparative studies focusing on flower synorganization and involving both families together are lacking.
For this study an extensive search through the morphological literature has been conducted. Based on this and my own studies on flowers in various Apocynaceae and Orchidaceae and complex flowers in other angiosperms with scanning electron microscopy and with microtome section series, a review on convergent floral traits in flower development and architecture in the two families is presented.
There is a tendency of protracted development of synorganized parts in Apocynaceae and Orchidaceae (development of synorganization of two or more organs begins earlier the more accentuated it is at anthesis). Synorganization (or complexity) also paves the way for novel structures. One of the most conspicuous such novel structures in Apocynaceae is the corona, which is not the product of synorganization of existing organs; however, it is probably enhanced by synorganization of other, existing, floral parts. In contrast to synorganized parts, the corona appears developmentally late.
Synorganization of floral organs may lead to a large number of convergences in clades that are only very distantly related. The convergences that have been highlighted in this comparative study should be developmentally investigated directly in parallel in future studies.
Journal Article
Flowers on the Tree of Life
Genetic and molecular studies have recently come to dominate botanical research at the expense of more traditional morphological approaches. This broad introduction to modern flower systematics demonstrates the great potential that floral morphology has to complement molecular data in phylogenetic and evolutionary investigations. Contributions from experts in floral morphology and evolution take the reader through examples of how flowers have diversified in a large variety of lineages of extant and fossil flowering plants. They explore angiosperm origins and the early evolution of flowers and analyse the significance of morphological characters for phylogenetic reconstructions on the tree of life. The importance of integrating morphology into modern botanical research is highlighted through case studies exploring specific plant groups where morphological investigations are having a major impact. Examples include the clarification of phylogenetic relationships and understanding the significance and evolution of specific floral characters, such as pollination mechanisms and stamen and carpel numbers.
eBook
The maleness of larger angiosperm flowers
Flower biomass varies widely across the angiosperms. Each plant species invests a given amount of biomass to construct its sex organs. A comparative understanding of how this limited resource is partitioned among primary (male and female structures) and secondary (petals and sepals) sexual organs on hermaphrodite species can shed light on general evolutionary processes behind flower evolution. Here, we use allometries relating different flower biomass components across species to test the existence of broad allocation patterns across the angiosperms. Based on a global dataset with flower biomass spanning five orders of magnitude, we show that heavier angiosperm flowers tend to be male-biased and invest strongly in petals to promote pollen export, while lighter flowers tend to be female-biased and invest more in sepals to insure their own seed set. This result demonstrates that larger flowers are not simple carbon copies of small ones, indicating that sexual selection via male–male competition is an important driver of flower biomass evolution and sex allocation strategies across angiosperms.
Journal Article
Reconstructing the ancestral angiosperm flower and its initial specializations
Increasingly robust understanding of angiosperm phylogeny allows more secure reconstruction of the flower in the most recent common ancestor of extant angiosperms and its early evolution. The surprising emergence of several extant and fossil taxa with simple flowers near the base of the angiosperms—Chloranthaceae, Ceratophyllum, Hydatellaceae, and the Early Cretaceous fossil Archaefructus (the last three are water plants)—has brought a new twist to this problem. We evaluate early floral evolution in angiosperms by parsimony optimization of morphological characters on phylogenetic trees derived from morphological and molecular data. Our analyses imply that Ceratophyllum may be related to Chloranthaceae, and Archaefructus to either Hydatellaceae or Ceratophyllum. Inferred ancestral features include more than two whorls (or series) of tepals and stamens, stamens with protruding adaxial or lateral pollen sacs, several free, ascidiate carpels closed by secretion, extended stigma, extragynoecial compitum, and one or several ventral pendent ovule(s). The ancestral state in other characters is equivocal: e.g., bisexual vs. unisexual flowers, whorled vs. spiral floral phyllotaxis, presence vs. absence of tepal differentiation, anatropous vs. orthotropous ovules. Our results indicate that the simple flowers of the newly recognized basal groups are reduced rather than primitively simple.
Journal Article
Evolution and genetic control of the floral ground plan
The floral ground plan is a map of where and when floral organ primordia arise. New results combining the defined phylogeny of flowering plants with extensive character mapping have predicted that the angiosperm ancestor had whorls rather than spirals of floral organs in large numbers, and was bisexual. More confidently, the monocot ancestor likely had three organs in each whorl, whereas the rosid and asterid ancestor (Pentapetalae) had five, with the perianth now divided into sepals and petals. Genetic mechanisms underlying the establishment of the floral ground plan are being deduced using model species, the rosid Arabidopsis, the asterid Antirrhinum, and in grasses such as rice. In this review, evolutionary and genetic conclusions are drawn together, especially considering how known genes may control individual processes in the development and evolution of ground plans. These components include organ phyllotaxis, boundary formation, organ identity, merism (the number or organs per whorl), variation in the form of primordia, organ fusion, intercalary growth, floral symmetry, determinacy and, finally, cases where the distinction between flowers and inflorescences is blurred. It seems likely that new pathways of ground plan evolution, and new signalling mechanisms, will soon be uncovered by integrating morphological and genetic approaches.
Journal Article
Evolution of flower allometry and pigmentation in Mammillaria haageana (Cactaceae)
Background
A puzzle in evolution is the understanding of how the environment might drive subtle phenotypic variation, and whether this variation is adaptive. Under the neutral evolutionary theory, subtle phenotypes are almost neutral with little adaptive value. To test this idea, we studied the infraspecific variation in flower shape and color in
Mammillaria haageana
, a species with a wide geographical distribution and phenotypic variation, which populations are often recognized as infraspecific taxa.
Results
We collected samples from wild populations, kept them in the greenhouse for at least one reproductive year, and collected newly formed flowers. Our first objective was to characterize tepal natural variation in
M. haageana
through geometric morphometric and multivariate pigmentation analyses. We used landmark-based morphometrics to quantify the trends of shape variation and tepal color-patterns in 20
M. haageana
accessions, belonging to five subspecies, plus 8
M. albilanata
accessions for comparison as the sister species. We obtained eight geometric morphometric traits for tepal shape and color-patterns. We found broad variation in these traits between accessions belonging to the same subspecies, without taxonomic congruence with those infraspecific units. Also the phenetic cluster analysis showed different grouping patterns among accessions. When we correlated these phenotypes to the environment, we also found that solar radiation might explain the variation in tepal shape and color, suggesting that subtle variation in flower phenotypes might be adaptive. Finally we present anatomical sections in
M. haageana
subsp.
san-angelensis
to propose some of the underlying tepal structural features that may give rise to tepal variation.
Conclusions
Our geometric morphometric approach of flower shape and color allowed us to identify the main trends of variation in each accession and putative subspecies, but also allowed us to correlate these variation to the environment, and propose anatomical mechanisms underlying this diversity of flower phenotypes.
Journal Article
The evolution of floral biology in basal angiosperms
In basal angiosperms (including ANITA grade, magnoliids, Choranthaceae, Ceratophyllaceae) almost all bisexual flowers are dichogamous (with male and female functions more or less separated in time), and nearly 100 per cent of those are protogynous (with female function before male function). Movements of floral parts and differential early abscission of stamens in the male phase are variously associated with protogyny. Evolution of synchronous dichogamy based on the day/night rhythm and anthesis lasting 2 days is common. In a few clades in Magnoliales and Laurales heterodichogamy has also evolved. Beetles, flies and thrips are the major pollinators, with various degrees of specialization up to large beetles and special flies in some large-flowered Nymphaeaceae, Magnoliaceae, Annonaceae and Aristolochiaceae. Unusual structural specializations are involved in floral biological adaptations (calyptras, inner staminodes, synandria and food bodies, and secretory structures on tepals, stamens and staminodes). Numerous specializations that are common in monocots and eudicots are absent in basal angiosperms. Several families are poorly known in their floral biology.
Journal Article
Flowers of the early-branching papilionoid legume Petaladenium urceoliferum display unique morphological and ontogenetic features
PREMISE OF THE STUDY: Floral development can help to shed light on puzzling features across flowering plants. The enigmatic Amazonian monospecific genus Petaladenium of the legume family (Leguminosae) had rarely been collected and only recently became available for ontogenetic studies. The fimbriate-glandular wing petals of P. urceoliferum are unique among the more than 19000 legume species. Ontogenetic data illuminate the systematic position of the genus and foster our understanding on floral evolution during the early diversification of the papilionoid legumes. METHODS: Flower buds were collected in the field, fixed in 70% ethanol, and investigated using scanning electron microscopy (SEM). Results were compared with existing material from early-diverging papilionoid legumes. KEY RESULTS: Formation of sepals and petals shows bidirectional tendencies. Stamens arise in two whorls, and the single carpel arises concomitantly with the outer stamen whorl. Gland formation starts early on the edges of the wing petals. The carpel reopens for a short time when the initiation of ovules is visible. Stornata at the base of the hypanthium indicate that the flower functions like other standard flag blossoms. CONCLUSIONS: The floral ontogeny confirms the close affinity of P. urceoliferum with the florally heterogeneous, early-diverging papilionoid Amburaneae clade. The results strengthen the theory of a distinct experimental phase among early-branching papilionoid legumes during which a wider range of floral morphologies arose. Polysymmetry, monosymmetry, variable organ numbers, and a wide range of ontogenetic patterns laid the foundation for a successful canalization toward the more restricted but well-adapted dorsiventral papilionoid flag blossom.
Journal Article