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Paula J Rudall

Publications and source records attributed to Paula J Rudall.

10 recordsLinked to original sources

Evolutionary and morphometric implications of morphological variation among flowers within an inflorescence: a case-study using European orchids.

BACKGROUND AND AIMS: This study explores the previously largely ignored morphological variation that occurs among flowers within a single inflorescence. METHODS: Variation in four metric parameters (labellum length and width, spur length and width) that together strongly influence pollination frequency is documented within the simple racemose inflorescences of eight individuals that represent a primary hybrid and six species of European orchids. KEY RESULTS: Regression of each parameter against the location of each flower on the inflorescence, and calculation of correlation coefficients for each pair of parameters within each inflorescence, demonstrate significant decoupling of labellum and spur development, despite the fact that they are different portions of the same floral organ. Spur length and diameter are constant across inflorescences of Dactylorhiza other than the vestigial-spurred D. viridis, whereas in other genera spur length declines in parallel with labellum dimensions. These differences are likely to reflect selection pressures or developmental constraints. Strong negative deviations from the regression line for one or more parameters are evident in occasional flowers, occurring most frequently in the lowermost and uppermost one or two flowers, and so reflecting transitions in meristematic behaviour. Thus, population-level morphometric studies are best conducted on flowers taken from approximately the mid-point of the inflorescence. Moreover, in the two relatively large inflorescences where lower flowers were removed for measurement before the upper flowers had opened, labellum size increased significantly in the flowers immediately above the excisions, suggesting that excision liberated resources that were diverted into the opening buds. Repeat measurement of all flowers from one selected inflorescence demonstrated typical measurement errors of only +/- 30-80 micro m, irrespective of the size of the structure studied. If flowers are not mounted and measured immediately following excision, modest negative deviations of 30-50 micro m result from post-mounting shrinkage; this occurs less rapidly in the spur than in the thinner labellum, which should therefore be measured first. Variation in all four parameters among all the flowers of a single inflorescence is between 42 % and 107 % of that observed between a similar number of flowers sampled from a consistent location on different (but conspecific and coexisting) inflorescences. CONCLUSIONS: This result demonstrates the strong influence of epigenesis on flower morphology and further emphasizes the importance of (a) sampling from a consistent location within the inflorescences under comparison, (b) interpreting morphometric ordinations hierarchically, building from individuals to infraspecific taxa and species via populations, and (c) considering in any microevolutionary study the potentially profound effects of the cline in flower size within each inflorescence.

Biological Evolution↗

Flower-like terminal structures in racemose inflorescences: a tool in morphogenetic and evolutionary research.

Terminal flower-like structures (TFLS) occur in many angiosperms that possess indeterminate inflorescences such as spikes, racemes, or spadices. We describe and review TFLS in early-divergent angiosperms, especially the magnoliid order Piperales and the monocot order Alismatales, in which floral interpretation is controversial. Essentially similar TFLS occur in a wide range of taxa. Among magnoliids, they occur in some Piperales (Saururaceae and a few Piperaceae), but are absent from Chloranthaceae. Among monocots, they occur in some early-divergent families such as Acoraceae, Aponogetonaceae, Juncaginaceae, Potamogetonaceae, and Ruppiaceae. Similar TFLS with obscure organ identity are recorded in mutants of Arabidopsis. TFLS can often be interpreted as pseudanthia (close aggregations of reduced flowers), but in some cases the entire terminal pseudanthium is very similar to a true flower. In some cases, elaborated TFLS could therefore have given rise to what are normally termed 'true' (i.e. euanthial) flowers. Data presented here on terminal pseudanthia in Potamogeton and Ruppia support a pseudanthial evolutionary origin of reproductive units in the alismatid families Zannichelliaceae and Cymodoceaceae. Furthermore, in some alismatid species, either the entire inflorescence apex or an individual primordium at or near the inflorescence tip can be transformed into a filamentous or tubular (or intermediate) structure. A tubular structure enclosing stamens and carpels is described in Piper. This indicates that pseudanthium formation can provoke morphological novelties, perhaps due to new patterns of overlap between expression zones of regulatory genes and/or new spatial constraints.

Alismatales↗

How many nuclei make an embryo sac in flowering plants?

Research on early-divergent angiosperms, including Amborella, the putative sister to all other extant angiosperms, is increasingly used as a yardstick to infer the nature of the hypothetical ancestral angiosperm. Some traits are relatively diverse (and hence relatively labile) in this phylogenetic grade, compared with the more derived eudicot clade, in which developmental patterns have become increasingly canalized. One of the many mysteries surrounding the origin of the angiosperms is the evolutionary origin of the Polygonum-type embryo sac (monosporic, eight-nucleate and seven-celled) that occurs in the majority of flowering plants. Observations on the megagametophyte of Amborella are conflicting, but a recent report of a supernumerary synergid in this genus raises the question of whether the Polygonum-type embryo sac is derived by duplication of a four-nucleate structure or by reduction from a multicellular structure.

Cell Nucleus↗

Morphological and molecular phylogenetic context of the angiosperms: contrasting the 'top-down' and 'bottom-up' approaches used to infer the likely characteristics of the first flowers.

Recent attempts to address the long-debated 'origin' of the angiosperms depend on a phylogenetic framework derived from a matrix of taxa versus characters; most assume that empirical rigour is proportional to the size of the matrix. Sequence-based genotypic approaches increase the number of characters (nucleotides and indels) in the matrix but are confined to the highly restricted spectrum of extant species, whereas morphology-based approaches increase the number of phylogenetically informative taxa (including fossils) at the expense of accessing only a restricted spectrum of phenotypic characters. The two approaches are currently delivering strongly contrasting hypotheses of relationship. Most molecular studies indicate that all extant gymnosperms form a natural group, suggesting surprisingly early divergence of the lineage that led to angiosperms, whereas morphology-only phylogenies indicate that a succession of (mostly extinct) gymnosperms preceded a later angiosperm origin. Causes of this conflict include: (i) the vast phenotypic and genotypic lacuna, largely reflecting pre-Cenozoic extinctions, that separates early-divergent living angiosperms from their closest relatives among the living gymnosperms; (ii) profound uncertainty regarding which (a) extant and (b) extinct angiosperms are most closely related to gymnosperms; and (iii) profound uncertainty regarding which (a) extant and (b) extinct gymnosperms are most closely related to angiosperms, and thus best serve as 'outgroups' dictating the perceived evolutionary polarity of character transitions among the early-divergent angiosperms. These factors still permit a remarkable range of contrasting, yet credible, hypotheses regarding the order of acquisition of the many phenotypic characters, reproductive and vegetative, that distinguish 'classic' angiospermy from 'classic' gymnospermy. The flower remains ill-defined and its mode (or modes) of origin remains hotly disputed; some definitions and hypotheses of evolutionary relationships preclude a role for the flower in delimiting the angiosperms. We advocate maintenance of parallel, reciprocally illuminating programmes of morphological and molecular phylogeny reconstruction, respectively supported by homology testing through additional taxa (especially fossils) and evolutionary-developmental genetic studies that explore genes potentially responsible for major phenotypic transitions.

Botany↗

How much data are needed to resolve a difficult phylogeny?: case study in Lamiales.

Reconstructing phylogeny is a crucial target of contemporary biology, now commonly approached through computerized analysis of genetic sequence data. In angiosperms, despite recent progress at the ordinal level, many relationships between families remain unclear. Here we take a case study from Lamiales, an angiosperm order in which interfamilial relationships have so far proved particularly problematic. We examine the effect of changing one factor-the quantity of sequence data analyzed-on phylogeny reconstruction in this group. We use simulation to estimate a priori the sequence data that would be needed to resolve an accurate, supported phylogeny of Lamiales. We investigate the effect of increasing the length of sequence data analyzed, the rate of substitution in the sequences used, and of combining gene partitions. This method could be a valuable technique for planning systematic investigations in other problematic groups. Our results suggest that increasing sequence length is a better way to improve support, resolution, and accuracy than employing sequences with a faster substitution rate. Indeed, the latter may in some cases have detrimental effects on phylogeny reconstruction. Further molecular sequencing-of at least 10,000 bp-should result in a fully resolved and supported phylogeny of Lamiales, but at present the problematic aspects of this tree model remain.

Base Sequence↗

Three-dimensional analysis of plant structure using high-resolution X-ray computed tomography.

High-resolution X-ray computed tomography (HRCT) is a non-invasive approach to 3D visualization and quantification of biological structure. The data, based on differential X-ray attenuation, are analogous to those otherwise obtainable only by serial sectioning. Requiring no fixing, sectioning or staining, HRCT produces a 3D digital map of the specimen that allows measurements and visualizations, including arbitrarily oriented sections. In spite of its application throughout the natural sciences, HRCT has yet to be applied in extant plant structural research.

Imaging, Three-Dimensional↗

Evolutionary change in flowers and inflorescences: evidence from naturally occurring terata.

Records of naturally occurring, heritable floral abnormalities considerably enhance our understanding of floral evolution. Peloric mutants, frequent in natural populations of orchids and mints, have radially symmetric flowers but occur in species characterized by bilaterally symmetric flowers. Three distributions of peloric flowers across an inflorescence are: (1) complete (all flowers peloric, as in the cycloidea mutant of Antirrhinum), (2) scattered (with both peloric and zygomorphic flowers, as in the epigenetic cycloidea mutant of Linaria), and (3) terminal (only the terminal flower peloric, as in the centroradialis mutant of Antirrhinum). Genetic relationships between lateral and terminal peloria, and between peloric and pseudopeloric flowers, remain ambiguous. Complete peloria probably caused occasional evolutionary reversals from zygomorphy to actinomorphy, whereas the 'terminal-flower effect' is a less likely cause of floral evolution.

Biological Evolution↗

Systematic significance of cell inclusions in Haemodoraceae and allied families: silica bodies and tapetal raphides.

This paper presents the first record of silica deposits in tissues of Haemodoraceae and adds new records of tapetal raphides in this family. Within the order Commelinales, silica is present in leaves of three families (Hanguanacaeae, Haemodoraceae and Commelinaceae), but entirely absent from the other two (Pontederiaceae and Philydraceae). Presence or absence of characteristic cell inclusions may have systematic potential in commelinid monocotyledons, although the existing topology indicates de novo gains and losses in individual families. Silica sand was observed in leaves of five out of nine genera examined of Haemodoraceae, predominantly in vascular bundle sheath cells and epidermal cells. Within Haemodoraceae, silica is limited to subfamily Conostylidoideae. The occurrence of silica in Phlebocarya supports an earlier transfer of this genus from Haemodoroideae to Conostylidoideae. The presence of raphides (calcium oxalate crystals) in the anther tapetum represents a rare character, only reported in a few monocot families of the order Commelinales, and possibly representing a mechanism for regulation of cytoplasmic free calcium levels. Tapetal raphides were observed here in Anigozanthus and Conostylis (both Haemodoraceae), and Tradescantia (Commelinaceae), thus supplementing two earlier records in Haemodoraceae, Philydraceae and Commelinaceae.

Calcium Oxalate↗

Roles of synorganisation, zygomorphy and heterotopy in floral evolution: the gynostemium and labellum of orchids and other lilioid monocots.

A gynostemium, comprising stamen filaments adnate to a syncarpous style, occurs in only threc groups of monocots: the large family Orchidaceae (Asparagales) and two small genera Pauridia (Hypoxidaceae: Asparagales) and Corsia (Corsiaceae, probably in Liliales), all epigynous taxa. Pauridia has actinomorphic (polysymmetric) flowers, whereas those of Corsia and most orchids are strongly zygomorphic (monosymmetric) with a well-differentiated labellum. In Corsia the labellum is formed from the outer median tepal (sepal), whereas in orchids it is formed from the inner median tepal (petal) and is developmentally adaxial (but positionally abaxial in orchids with resupinate flowers). Furthermore, in orchids zygomorphy is also expressed in the stamen whorls, in contrast to Corsia. In Pauridia a complete stamen whorl is suppressed, but the 'lost' outer whorl is fused to the style. The evolution of adnation and zygomorphy are discussed in the context of the existing phylogenetic framework in monocotyledons. An arguably typological classification of floral terata is presented, focusing on three contrasting modes each of peloria and pseudopeloria. Dynamic evolutionary transitions in floral morphology are assigned to recently revised concepts of heterotopy (including homeosis) and heterochrony, seeking patterns that delimit developmental constraints and allow inferences regarding underlying genetic controls. Current evidence suggests that lateral heterotopy is more frequent than acropetal heterotopy, and that full basipetal heterotopy does not occur. Pseudopeloria is more likely to generate a radically altered yet functional perianth, but is also more likely to cause acropetal modification of the gynostemium. These comparisons indicate that there are at least two key genes or sets of genes controlling adnation, adaxial stamen suppression and labellum development in lilioid monocots; at least one is responsible for stamen adnation to the style (i.e. gynostemium formation), and another controls adaxial stamen suppression and adaxial labellum formation in orchids. Stamen adnation to the style may be a product of over-expression of the genes related to epigyny (i.e. a form of hyper-epigyny). If, as seems likely, stamen-style adnation preceded zygomorphy in orchid evolution, then the flowers of Pauridia may closely resemble those of the immediate ancestors of Orchidaceae, although existing molecular phylogenetic data indicate that a sister-group relationship is unlikely. The initial radiation in Orchidaceae can be attributed to the combination of hyper-epigyny, zygomorphy and resupination, but later radiations at lower taxonomic levels that generated the remarkable species richness of subfamilies Orchidoideae and Epidendroideae are more likely to reflect more subtle innovations that directly influence pollinator specificity, such as the development of stalked pollinaria and heavily marked and/or spur-bearing labella.

Biological Evolution↗