Paula J. Rudall

Systematics and Biology of Silica Bodies in Monocotyledons

Many plants take up soluble monosilicic acid from the soil. Some of these plants subsequently deposit it as cell inclusions of characteristic structure. This article describes the distribution and diversity of opaline silica bodies in monocotyledons in a phylogenetic framework, together with a review of techniques used for their examination, and the ecology, function and economic applications of these cell inclusions. There are several different morphological forms of silica in monocot tissues, and the number of silica bodies per cell may also vary. The most common type is the “druse-like” spherical body, of which there is normally a single body per cell, more in some cases. Other forms include the conical type and an amorphous, fragmentary type (silica sand). Silica bodies are most commonly found either in the epidermis (e.g., in grasses, commelinas and sedges) or in the sheath cells of vascular bundles (e.g., in palms, bananas and orchids). Silica-bearing cells are most commonly associated either with subepidermal sclerenchyma or bundle-sheath sclerenchyma. Silica bodies are found only in orchids and commelinids, not in other lilioid or basal monocots. In orchids, silica bodies are entirely absent from subfamilies Vanilloideae and Orchidoideae and most Epidendroideae but present in some Cypripedioideae and in the putatively basal orchid subfamily Apostasioideae. Among commelinid monocots, silica bodies are present in all palms, Dasypogonaceae and Zingiberales but present or absent in different taxa of Poales and Commelinales, with at least four separate losses of silica bodies in Poales.

Pointillist structural color in Pollia fruit

Biological communication by means of structural color has existed for at least 500 million years. Structural color is commonly observed in the animal kingdom, but has been little studied in plants. We present a striking example of multilayer-based strong iridescent coloration in plants, in the fruit of Pollia condensata. The color is caused by Bragg reflection of helicoidally stacked cellulose microfibrils that form multilayers in the cell walls of the epicarp. We demonstrate that animals and plants have convergently evolved multilayer-based photonic structures to generate colors using entirely distinct materials. The bright blue coloration of this fruit is more intense than that of any previously described biological material. Uniquely in nature, the reflected color differs from cell to cell, as the layer thicknesses in the multilayer stack vary, giving the fruit a striking pixelated or pointillist appearance. Because the multilayers form with both helicoidicities, optical characterization reveals that the reflected light from every epidermal cell is polarized circularly either to the left or to the right, a feature that has never previously been observed in a single tissue.

Molecular systematics of Iridaceae : evidence from four plastid DNA regions

Iridaceae are one of the largest families of Lilianae and probably also among the best studied of monocotyledons. To further evaluate generic, tribal, and subfamilial relationships we have produced four plastid DNA data sets for 57 genera of Iridaceae plus outgroups: rps4, rbcL (both protein-coding genes), the trnL intron, and the trnL-F intergenic spacer. All four matrices produce similar although not identical trees, and we thus analyzed them in a combined analysis, which produced a highly resolved and well-supported topology, in spite of the fact that the partition homogeneity test indicated strong incongruence. In each of the individual trees, some genera or groups of genera are misplaced relative to morphological cladistic studies, but the combined analysis produced a pattern much more similar to these previous ideas of relationships. In the combined tree, all subfamilies were resolved as monophyletic, except Nivenioideae that formed a grade in which Ixioideae were embedded. Achlorophyllous Geosiris (sometimes referred to Geosiridaceae or Burmanniaceae) fell within the nivenioid grade. Most of the tribes were monophyletic, and Isophysis (Tasmanian) was sister to the rest of the family; Diplarrhena (Australian) fell in a well-supported position as sister to Irideae/Sisyrinchieae/Tigridieae/Mariceae (i.e., Iridoideae); Bobartia of Sisyrinchieae is supported as a member of Irideae. The paraphyly of Nivenioideae is suspicious due to extremely high levels of sequence divergence, and when they were constrained to be monophyletic the resulting trees were only slightly less parsimonious (<1.0%). However, this subfamily also lacks clear morphological synapomorphies and is highly heterogeneous, so it is difficult to develop a strong case on nonmolecular grounds for their monophyly.

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 three 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.

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.

Evolution of Microsporogenesis in Angiosperms

Microsporogenesis is highly labile in early‐branching angiosperms, i.e., those with mostly sulcate pollen, compared with the tricolpate and tricolpate‐derived eudicots. New records of microsporogenesis in basal angiosperms (19 taxa were examined), together with a review of the literature, demonstrate that the existing typology has been too strictly applied; several basal angiosperms have apparently intermediate forms and therefore do not fit easily into simultaneous or successive categories. Intermediate forms include the “modified simultaneous” type, where ephemeral cell plates are formed after the first meiotic division but then disperse, and simultaneous cleavage follows the second meiotic division. This relative diversity reflects a range of variation in number and position of pollen apertures in basal angiosperms, although both monosulcate and inaperturate pollen may occur in conjunction with either simultaneous or successive microsporogenesis. However, many taxa with inaperturate pollen have successive microsporogenesis, whereas many monosulcate taxa have the simultaneous type (although successive and monosulcate is common in monocotyledons). The predominance of simultaneous microsporogenesis in extant basal angiosperms and in land plants in general (including gymnosperms) indicates that simultaneous microsporogenesis is plesiomorphic in angiosperms, despite the occurrence of the successive type in the putative first‐branching extant angiosperm, Amborella. This conclusion contradicts earlier views on the evolutionary polarity of this character.

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.

INAPERTURATE POLLEN IN MONOCOTYLEDONS

Inaperturate pollen is widespread in the monocotyledons, includes a diverse range of forms, and has arisen independently numerous times. Evidence for this comes from the phylogenetic distribution of inaperturate pollen and also from developmental and structural differences. There is no correlation between the production of inaperturate grains and either microsporogenesis type or tapetum type. Inaperturate pollen has phylogenetic significance within some groups, e.g., in Zingiberales, Liliales, and some Asparagales. Character states for inaperturate pollen are presented. It can broadly be divided into two types, “omniaperturate” and “functionally monoaperturate,” based on the thickening of the intine. The characters of omniaperturate pollen are adaptations that potentially increase the germination efficiency of the pollen. Both types sometimes occur in environments where pollen is not subject to desiccation and, thus, may remain viable with reduced exines.

Yams reclassified: a recircumscription of Dioscoreaceae and Dioscoreales

Analyses of morphological and molecular characters for Dioscoreales Hook, f. (Chase & al., 1995b; Caddick & al., 2000a; Caddick & al., 2002) have redefined the order, which now comprises three families, Burmanniaceae, Dioscoreaceae, and Nartheciaceae. Since recent analyses of morphological and molecular data sets (Caddick & al., 2002) have indicated well-supported relationships within Dioscoreaceae R. Br., a formal reclassification of the family is presented here. Dioscoreaceae now contain four distinct genera, Dioscorea, Stenomeris, Tacca (previously in Taccaceae), and Trichopus. The Malagasy endemic Avetra sempervirens is close sister to Trichopus zeylanicus, and is here reclassified as a second species of this genus. The dioecious Dioscoreaceae genera, Borderea, Epipetrum, Nanarepenta, Rajania, Tamus, and Testudinaria, are nested within Dioscorea in phylogenetic analyses (Caddick & al., 2002), and are therefore sunk into it.

Evolution of reproductive structures in grasses (Poaceae) inferred by sister-group comparison with their putative closest living relatives, Ecdeiocoleaceae.

Despite much recent activity in the phylogeny and developmental genetics of grasses, the enigmatic homologies of their reproductive structures remain largely unresolved, partly because their highly derived morphology has resulted in a unique associated terminology. Outstanding questions include whether grass lodicules and stamens are derived from a single perianth or stamen whorl, respectively, whether the grass caryopsis is homologous with a nut, and how the scutellum evolved. We investigated the reproductive structures of the putative sister group of grasses, the southwestern Australian family Ecdeiocoleaceae, which includes two genera, Ecdeiocolea and Georgeantha. The zygomorphic arrangement of the four (rather than six) stamens in male flowers of Ecdeiocolea indicates that they may represent three outer stamens plus the adaxial inner stamen. Within Ecdeiocoleaceae, characters such as the highly unusual nucellus structure of Ecdeiocolea are autapomorphic. Sister-group comparisons indicate that some characteristic grass features, notably the scutellum, do not occur in their putative closest relatives and that more data are needed on early-diverging grass genera to resolve these issues. The grass caryopsis could represent one end of a transformation series embodied by the reduced gynoecial structure and indehiscent fruit of other Poales such as Flagellaria, Joinvillea, and Ecdeiocolea.