Floral Organ Initiation Research Articles

A brassinolide‐suppressed rice MADS‐box transcription factor, OsMDP1, has a negative regulatory role in BR signaling

The MADS-box transcription factor-encoding genes are expressed mainly during plant reproductive development, where they play important roles in controlling floral organ initiation and identity. Few previous reports have investigated the functions of MADS-box transcription factors expressed in vegetative tissues. Here we describe the functional characterization of a rice AG-like MADS-box protein, OsMDP1 (Oryza sativa MADS-domain-containing protein 1). A partial cDNA encoding a MADS-box domain was identified via high-throughput screening of rice brassinolide-regulated genes, and the full-length cDNA was subsequently isolated via screening of a cDNA library constructed from rice materials at tillering stage. Expression pattern analyses indicated that OsMDP1 is transcribed mainly in vegetative tissues, including the mature leaf, coleoptile, root-elongation zone, culm internode, and especially the joint region between the leaf blade and sheath. Further studies revealed that transcription of OsMDP1 is stimulated by darkness and suppressed by brassinolide treatment. OsMDP1 deficiency resulted in shortened primary roots, elongated coleoptiles and enhanced lamina joint inclinations. Moreover, transgenic plants showed hypersensitivities to exogenous brassinolide in terms of lamina joint inclination and coleoptile elongation. OsMDP1 deficiency resulted in enhanced expression of OsXTR1, which encodes xyloglucan endotransglycosylase, the cell-wall loosening enzyme necessary for cell elongation, and modulated expressions of multiple genes involved in cell signalling and gene transcription, indicating the key negative regulatory role of OsMDP1 in BR signalling.

Comparative floral anatomy and ontogeny in Magnoliaceae

Floral anatomy and ontogeny are described in six species of Magnoliaceae, representing the two subfamilies Liriodendroideae (Liriodendron chinese and L. tulipifera) and Magnolioideae, including species with terminal flowers (Magnolia championi, M. delavayi, M. grandiflora, M. paenetalauma) and axillary flowers (Michelia crassipes). The sequence of initiation of floral organs is from proximal to distal. The three distinct outermost organs are initiated in sequence, but ultimately form a single whorl; thus their ontogeny is consistent with a tepal interpretation. Tepals are initiated in whorls, and the stamens and carpels are spirally arranged, though the androecium shows some intermediacy between a spiral and whorled arrangement. Carpels are entirely free from each other both at primordial stages and maturity. Ventral closure of the style ranges from open in Magnolia species examined to partially closed in Michelia crassipes and completely closed in Liriodendron, resulting in a reduced stigma surface. Thick-walled cells and tannins are present in all species except Michelia crassipes. Oil cells are normally present. Floral structure is relatively homogeneous in this family, although Liriodendron differs from other Magnoliaceae in that the carpels are entirely closed at maturity, resulting in a relatively small stigma, in contrast to the elongate stigma of most species of Magnolia. The flower of Magnolia does not terminate in an organ or organ whorl but achieves determinacy by gradual diminution.

Floral and Vegetative Morphogenesis in California Poppy (Eschscholzia californicaCham.)

For studies of the evolution of development in angiosperms, early‐diverging eudicot taxa are of particular interest for comparisons with established core eudicot model plants, such as Arabidopsis. Here we provide a detailed description of shoot and floral development of the basal eudicot California poppy (Eschscholzia californica). Rosette formation in the vegetative phase is accompanied by increased leaf complexity and shoot apex size. The flowering phase is characterized by internode elongation, formation of dissected cauline leaves, terminal flowers, and basipetal inflorescence branching. For developing flowers and fruits, we have defined 14 stages according to important landmark events, from inflorescence primordium initiation through seed dispersal. Floral organ initiation, morphogenesis, increase in floral meristem size, and the surface structure of mature floral organs are recorded in detail. The duration of the later floral stages, as well as the path of pollen tube growth in the gynoecium, is documented. Comparison of California poppy floral development with that of Arabidopsis indicates considerable differences in terms of organ fusion, whorl proliferation, and variability of size and organ number between the two species. Transitions in meristem identity from germination to floral organogenesis were monitored using expression of the developmental control gene EcFLO, the Eschscholzia ortholog of FLORICAULA/LEAFY. We found that the pattern of expression of EcFLO in the flanks of the shoot apex is maintained from late embryogenesis to flower initiation, indicating a continuous role for this gene in meristem function. As flower organs develop, EcFLO expression becomes more restricted to petal and stamen primordia. Development of the gynoecium occurs without EcFLO expression, indicating that EcFLO may not be necessary for the activation of C‐class genes.

Floral Patterning in Lotus japonicus

Floral patterning in Papilionoideae plants, such as pea (Pisum sativum) and Medicago truncatula, is unique in terms of floral organ number, arrangement, and initiation timing as compared to other well-studied eudicots. To investigate the molecular mechanisms involved in the floral patterning in legumes, we have analyzed two mutants, proliferating floral meristem and proliferating floral organ-2 (pfo-2), obtained by ethyl methanesulfonate mutagenesis of Lotus japonicus. These two mutants showed similar phenotypes, with indeterminate floral structures and altered floral organ identities. We have demonstrated that loss of function of LjLFY and LjUFO/Pfo is likely to be responsible for these mutant phenotypes, respectively. To dissect the regulatory network controlling the floral patterning, we cloned homologs of the ABC function genes, which control floral organ identity in Arabidopsis (Arabidopsis thaliana). We found that some of the B and C function genes were duplicated. RNA in situ hybridization showed that the C function genes were expressed transiently in the carpel, continuously in stamens, and showed complementarity with the A function genes in the heterogeneous whorl. In proliferating floral meristem and pfo-2 mutants, all B function genes were down-regulated and the expression patterns of the A and C function genes were drastically altered. We conclude that LjLFY and LjUFO/Pfo are required for the activation of B function genes and function together in the recruitment and determination of petals and stamens. Our findings suggest that gene duplication, change in expression pattern, gain or loss of functional domains, and alteration of key gene functions all contribute to the divergence of floral patterning in L. japonicus.

Transformation of maize with the p1 transcription factor directs production of silk maysin, a corn earworm resistance factor, in concordance with a hierarchy of floral organ pigmentation

The maize p1 gene encodes an R2R3-MYB transcription factor that controls the biosynthesis of red flavonoid pigments in floral tissues of the maize plant. Genetic and quantitative trait locus analyses have also associated the p1 gene with the synthesis of maysin, a flavone glycoside from maize silks that confers natural resistance to corn earworm. Here, we show directly that the p1 gene induces maysin accumulation in silk tissues. Transformation of maize plants that had low or no silk maysin with p1 transgenes elevated silk maysin concentrations to levels sufficient for corn earworm abiosis. The p1 transgenes also conferred red pigment to pericarp, cob, husk and tassel tissues, as expected; however, different subsets of these tissues were pigmented within individual transgenic plants. Statistical analysis shows that the pigmentation patterns observed amongst the p1 transgenic plants conform to a hierarchy that is similar to the temporal ordering of floral organ initiation. We propose that the observed hierarchy of pigmentation patterns is conferred by variation due to epigenetic control of the p1 transgenes. The production of plants with improved traits through genetic engineering can depend in large part on the achievement of tight organ-specific expression of the introduced transgenes. Our results demonstrate that the production of transgenic plants using a promoter with well-defined tissue specificity, such as the p1 promoter, can result in unexpected variation in tissue specificity amongst the resulting transgenic plants.

The role of SEUSS in auxin response and floral organ patterning.

Genetic and physiological analyses implicate auxin flux in patterning, initiation and growth of floral organs. Within the Arabidopsis flower, the ETTIN/ARF3 transcription factor responds to auxin to effect perianth organ number and reproductive organ differentiation. This work describes a modifier of ettin that causes filamentous, mispositioned outer whorl organs and reduced numbers of malformed stamens in the double mutant. The modifier was discovered to be a new allele of the seuss (seu) mutant. SEU encodes a novel protein that is predicted to transcriptionally co-repress the AGAMOUS floral organ identity gene. The effects of seu on ett are shown to be independent of the SEU-AG pathway. Furthermore, morphological, physiological and genetic evidence implicate SEU in auxin-regulated growth and development. seu has a pleiotropic phenotype that includes reductions in several classic auxin responses such as apical dominance, lateral root initiation, sensitivity to exogenous auxin and activation of the DR5 auxin response reporter. seu displays a synergistic interaction with the auxin response mutant pinoid, producing flowers with few outer whorl organs. Collectively, these data suggest that SEU is a novel factor affecting auxin response. A model is proposed in which SEU functions jointly with ETT in auxin response to promote floral organ patterning and growth.

Floral development in Polygala myrtifolia (Polygalaceae) and its similarities with Leguminosae

Floral development of Polygala myrtifolia was studied by the use of SEM, and compared with floral development in Leguminosae, with which Polygalaceae is grouped in Fabales. Five sepals and petals are formed in a helix, the direction of which is predetermined by the sequentially formed bracteoles. The two lateral sepals are delayed in their development, and the two lateral petals are suppressed early. For stamen initiation a hypothetical pathway is presented, in which the initiation is in two helical whorls. The abaxial, antesepalous stamen and the adaxial, antepetalous stamen are lost. In the gynoecium the late closure is distinct. Although the floral morphology of Polygalaceae and Leguminosae differs clearly, the two families do share some developmental features: (1) The formation of a racemose inflorescence, (2) the acropetal and helical initiation of floral organs, (3) the morphogenetic function of bracteoles, (4) the suppression and loss of organs, (5) the synchronous stamen differentiation (after asynchronous initiation) and late fusion of filaments, (6) the late closure of the gynoecium, and (7) the late expression of floral asymmetry. On the basis of these developmental similarities, the placement of Polygalaceae in Fabales becomes more plausible. Further insights in floral evolution of Fabales will be possible by means of a detailed phylogenetic analysis of developmental characters, based on a broadened data matrix.

Inflorescence development in an Ornithogalum dubium hybrid as influenced by bulb temperature treatments

SummaryOrnithogalum dubium hybrid 327 clone 2 (‘327-2’) bulbs were stored dry at 10°, 16°, 22°, 28°, and 35°C for six weeks after harvest. After storage, bulbs were evaluated by nuclear magnetic resonance imaging (MRI) to obtain the spin-lattice relaxation time (T1) profile across the transverse section of intact bulbs, by scanning electron microscopy (SEM) to observe inflorescence development, and by forcing in a 18.5°/18°C greenhouse to observe growth and flowering responses. Bulbs treated at 10°C had the shortest T1 (0.33 ms) through the bulb which is largely composed of scales. Leaf emergence from bulbs treated at 10°C was delayed, and plants failed to flower. This indicated that bulbs were dormant and dormancy was not broken, thus delaying initiation of floral organs. Bulbs treated at 22°C and 28°C formed the primary inflorescence with several florets. At the base of the primary scape of bulbs treated at 22°C, a vegetative apex was observed by both MRI and SEM. In the centre of bulbs where both leaves and floral organs were present, T1 was longer than that of the scales. This suggests that dormancy in the scales was broken and the leaves and scape were ready to emerge. Flowering was fastest when bulbs were treated at 22°C. The number of florets was the highest (16.0 florets) and fewest (12.6 florets) when bulbs were treated at 22°C and 35°C, respectively. Ornithogalum dubium bulbs stored at 25° and 30°C for six weeks flowered from 14 and 17 bulbs, respectively, out of 22 and flowered earlier than when stored either at lower or high temperatures. The response obtained O. dubium ‘327-2’ hybrid was attributed to its pedigree involving O. dubium. Due to its non-destructive nature, MRI can be used to observe inflorescence development inside the bulbs during bulb storage and possibly to study the state of dormancy.

Floral development of the model legume Medicago truncatula: ontogeny studies as a tool to better characterize homeotic mutations

This work provides new evidence of the complex genetic regulation necessary to accomplish flower development in legumes. Using scanning electron microscopy (SEM) analysis, we have characterized the early developmental events of the wild type Medicago truncatula flower and selected morphological characters as markers to break it down into eight different developmental stages. The order of floral organ initiation in M. truncatula and pea (Pisum sativum L.), in contrast to Arabidopsis and Antirrhinum, is unidirectional in all whorls starting from the abaxial position of the flower with a high degree of overlap. Another main difference is the existence of four common primordia from which petals and stamens differentiate. The formation of common primordia, as opposed to discrete petal and stamen primordia, has been described in many legume and non-legume plants. The main differences between pea and M. truncatula floral ontogeny are in carpel and fruit development. We also used these morphological markers as tools to characterize early alterations in the flower development of a male-sterile M. truncatula floral homeotic mutant named mtapetala. This mutant displays a phenotype resembling those of weak class B mutants with homeotic conversions of floral organ whorls 2 and 3 into sepaloid and carpelloid structures, respectively. Ontogeny studies of the mtapetala mutant flowers showed similarities with the effects of previously described loss-of-B-function mutations. Differences between ontogeny of wild type and mtapetala flowers could not be detected during the first stages (1–5) of flower development. In late stage 5, abnormal-shaped petals with acute lobes and trichomes as well as abnormal-shaped stamens were visible in whorls 2 and 3. At stage 6, the morphology of petals began to change, developing enlarged sepaloid structures bearing trichomes and first the antesepalous stamens and then the antepetalous stamens began to differentiate carpelloid anthers from filaments. Third whorl organs presented different degrees of carpelloidy. The present study should provide tools for the characterization and comparative analyses of new Medicago floral homeotic mutants and could be useful in elucidating how floral organ identity functions work in legumes.

FLOOZY of petunia is a flavin mono-oxygenase-like protein required for the specification of leaf and flower architecture.

The mechanisms that determine the relative positions of floral organs, and thereby their numbers, is a poorly understood aspect of flower development. We isolated a petunia mutant, floozy (fzy), in which the formation of floral organ primordia in the outermost three floral whorls and one of the two bracts at the base of the flower is blocked at an early stage. In addition, fzy mutants fail to generate secondary veins in leaves and bracts and display a decreased apical dominance in the inflorescence. FZY encodes an enzyme with homology to flavin mono-oxygenases and appears to be the ortholog of YUCCA genes of Arabidopsis. FZY is expressed in young leafs and bracts and in developing flowers. In young floral meristems FZY is expressed in the center of the meristem dome and, later, expression becomes localized on the flanks of the initiating petal and stamen primordia and at several sites in maturing anthers and carpels. These findings indicate that FZY is involved in synthesizing a signaling compound that is required for floral organ initiation and specification of the vascularization pattern in leaves. Although fzy mutants contain normal auxin levels, ectopic expression of FZY results in excessive auxin accumulation, suggesting that the signaling compound is auxin.

Floral organogenesis and development of two taxa of the Solanaceae-- Anisodus tanguticus and Atropa belladonna

Floral organogenesis and development of two Solanaceae species, Anisodus tanguticus and Atropa belladonna, were studied by using scanning electron microscopy (SEM) as part of a project on systematics and evolution in the tribe Hyoscyameae. These two species share the following common characters of floral organ initiation and development: (1) initiation of the floral organs in the two species follows Hofmeister's rule; (2) the mode of corolla tube development belongs to the late sympetaly type, namely, petals are initiated separately and later become joined by fusion of their basal meristem, then rise together and form a corolla tube; (3) primordia of the floral appendages are initiated in a pentamerous pattern and acropetal order: sepals are initiated first, followed by the petals and stamens, and finally the carpels. The whorl of five stamen primordia forms almost simultaneously and originates opposite the sepal primordia, but initiation of the sepal primordia shows different modes in the two species. The sepal primordia of Anisodus tanguticus have simultaneously whorled initiation, while those of Atropa belladonna have helical initiation. The systematic significance of the present results in the genera Anisodus and Atropa is discussed in this paper.

AINTEGUMENTA promotes petal identity and acts as a negative regulator of AGAMOUS.

The Arabidopsis AINTEGUMENTA (ANT) gene has been shown previously to be involved in ovule development and in the initiation and growth of floral organs. Here, we show that ANT acts in additional processes during flower development, including repression of AGAMOUS (AG) in second whorl cells, promotion of petal epidermal cell identity, and gynoecium development. Analyses of ap2-1 ant-6 double mutants reveal that ANT acts redundantly with AP2 to repress AG in second whorl cells. The abaxial surface of ant petals contains features such as stomata and elongated, interdigitated cells that are not present on wild-type petals. The loss of petal identity in these second whorl cells does not result from ectopic AG expression, suggesting that ANT acts in a pathway promoting petal cell identity that is independent of its role in repression of AG. These data suggest that ANT may function as a class A gene.

Ectopic expression of AINTEGUMENTA in Arabidopsis plants results in increased growth of floral organs.

AINTEGUMENTA (ANT) was previously shown to be involved in floral organ initiation and growth in Arabidopsis. ant flowers have fewer and smaller floral organs and possess ovules that lack integuments and a functional embryo sac. The present work shows that young floral meristems of ant plants are smaller than those in wild type. Failure to initiate the full number of organ primordia in ant flowers may result from insufficient numbers of meristematic cells. The decreased size of ant floral organs appears to be a consequence of decreased cell division within organ primordia. Ectopic expression of ANT under the control of the constitutive 35S promoter results in the development of larger floral organs. The number and shape of these organs is not altered and the size of vegetative organs is normal. Microscopic and molecular analyses indicate that the increased size of 35S::ANT sepals is the result of increased cell division, whereas the increased sizes of 35S::ANT petals, stamens, and carpels are primarily attributable to increased cell expansion. In addition, 35S::ANT ovules often exhibit increased growth of the nucellus and the funiculus. These results suggest that ANT stimulates cell growth in floral organs.

Control of floral homeotic gene expression and organ morphogenesis in Antirrhinum.

The development of reproductive organs in Antirrhinum depends on the expression of an organ identity gene, plena, in the central domain of the floral meristem. To investigate the mechanism by which plena is regulated, we have characterised three mutants in which the pattern of plena expression is altered. In polypetala mutants, expression of plena is greatly reduced, resulting in a proliferation of petals in place of reproductive organs. In addition, polypetala mutants exhibit an altered pattern of floral organ initiation, quite unlike that seen in loss-of-function plena mutants. This suggests that polypetala normally has two roles in flower development: regulation of plena and control of organ primordia formation. In fistulata mutants, plena is ectopically expressed in the distal domain of petal primordia, resulting in the production of anther-like tissue in place of petal lobes. Flowers of fistulata mutants also show a reduced rate of petal lobe growth, even in a plena mutant background. This implies that fistulata normally has two roles in the distal domain of petal primordia: inhibition of plena expression and promotion of lobe growth. A weak allele of the floral meristem identity gene, floricaula, greatly enhances the effect of fistulata on plena expression, showing that floricaula also plays a role in repression of plena in outer whorls. Taken together, these results show that genes involved in plena regulation have additional roles in the formation of organs, perhaps reflecting underlying mechanisms for coupling homeotic gene expression to morphogenesis.

Floral Ontogeny of Pelargonium × domesticum

Floral ontogeny of Pelargonium xdomesticum was examined for two cultivars grown under identical environmental conditions. Apical meristems of `Duchess' and `Jennifer' were vegetative at the commencement of the experiment. Meristems were examined every five days over an experimental period lasting 85 days. Floral ontogeny was the same for both cultivars, although the timing of floral initiation of the meristem, floral organ initiation, and floral organ development differed. The vegetative meristem was convex with leaf primordia initiated on either side in an alternate pattern. Early floral initiation was characterized by formation of a cleft towards one side of the meristem, followed quickly by a second cleft on the other side. Between the clefts new meristems developed. New meristems lacked leaf primordia. Proliferation of meristems continued until numerous meristems were organized in a cluster arrangement at the apex of the shoot. Proliferation of meristems at the apex continued until multiple inflorescences had developed. Inflorescences were subtended by bracts. Floral organ primordia were initiated in a succession of four whorls: sepals, petals, androecia and gynoecium. Petals and androecia appeared to develop simultaneously soon after sepal primordia were visible. Petal primordia remained small while the androecia continued to grow. The gynoecium first formed a conical shape, with carpels protruding from the base in a bulbous fashion. At the distal end of the gynoecium, divisions appeared which developed into stigmatic lobes. As the gynoecium elongated, stigmatic lobes became more pronounced. Petal elongation concurred in synchrony with elongation of the gynoecium. On one experimental unit of `Duchess' two florets had opened at 750 total cumulative moles. No `Jennifer' florets had opened by termination of the experiment.

Floral homeotic gene expression defines developmental arrest stages in Brassica oleracea L. vars. botrytis and italica.

Brassica oleracea L. vars, italica (broccoli) and botrytis (cauliflower) both undergo developmental arrests which result in heading phenotypes. We characterized these arrested tissues at the morphological and molecular levels, and defined the developmental changes that ensure after arrest has been broken. We found that the order of floral organ initiation and the positions of resulting floral organ primordia in this species in some respects from that of Arabidopsis, which is a member of the same family, Brassicaceae. We also cloned homologs of the Arabidopsis floral homeotic genes APETALA1 (AP1) and APETALA3 (AP3) from B. oleracea and characterized their expression patterns. We found that the AP1 homolog was expressed in some of the meristems of arrest-stage cauliflower, providing evidence that this tissue is florally determined. In broccoli, both the AP1 and AP3 homologs were expressed. However, the spatial pattern of expression of the broccoli AP1 homolog differed from that of Arabidopsis. In addition we identified a homolog of the CAULIFLOWER (CAL) gene, BoiCAL, from broccoli. The predicted amino acid sequence indicated that the BoiCAL gene product does not contain the mutation thought to be responsible for the early arrest exhibited in cauliflower (Kempin et al. 1995), but contains other changes that might play a role in the broccoli heading phenotype.

Trends in Evolution of Floral Ontogeny in Cassia Sensu Stricto, Senna, and Chamaecrista (Leguminosae: Caesalpinioideae: Cassieae: Cassiinae); A Study in Convergence

Distinctions in floral ontogeny among three segregate genera (Cassia sensu stricto, Chamaecrista, and Senna) of Cassia L. support their separation. In all species studied, the order of floral organ initiation is: sepals, petals, antesepalous stamens plus carpel, and lastly antepetalous stamens. Sepal initiation is helical in all three genera, which however differ in whether the first sepal is initiated in median abaxial position (Senna), or abaxial and off-median (Cassia javanica), a rare character state among legumes. Order of petal initiation varies: helical in Senna vs. unidirectional in Cassia and Chamaecrista. Both stamen whorls are uniformly unidirectional. Intergeneric ontogenetic differences occur in phyllotaxy, inflorescence architecture, bracteole formation, overlap of initiation among organ whorls (calyx/corolla in Cassia; two stamen whorls in Chamaecrista), eccentric initiation on one side of a flower, anther attachment, anther pore structure, and precocious carpel initiation in Senna. The asymmetric corolla and androecium in Chamaecrista arise by precocious organ initiation on one side (left or right). The poricidal anther character can result from differing developmental pathways: lateral slits vs. sealing of lateral sutures; clasping hairs vs. sutural ridges; terminal pores (one or two) vs. none; and clamp layer formation internally that prevents lateral dehiscence. Genera differ in corolla aestivation patterns and in stigma type. Convergence is shown among the three genera, based on intergeneric dissimilarities in early floral ontogeny (floral position in the inflorescence, bracteole presence, position of the first sepal initiated, order of petal initiation, asymmetric initiation, overlap between whorls, anther morphology, and time of carpel initiation) resulting in similarities at anthesis (showy, mostly yellow salverform flowers, heteromorphic stamens, poricidal anther dehiscence, bee pollination, and chambered stigma).

Trends in evolution of floral ontogeny in Cassia sensu stricto, Senna, and Chamaecrista (Leguminosae: Caesalpinioideae: Cassieae: Cassiinae); a study in convergence

Distinctions in floral ontogeny among three segregate genera (Cassia sensu stricto, Chamaecrista, and Senna) of Cassia L. support their separation. In all species studied, the order of floral organ initiation is: sepals, petals, antesepalous stamens plus carpel, and lastly antepetalous stamens. Sepal initiation is helical in all three genera, which however differ in whether the first sepal is initiated in median abaxial position (Senna), or abaxial and off‐median (Cassia javanica), a rare character state among legumes. Order of petal initiation varies: helical in Senna vs. unidirectional in Cassia and Chamaecrista. Both stamen whorls are uniformly unidirectional. Intergeneric ontogenetic differences occur in phyllotaxy, inflorescence architecture, bracteole formation, overlap of initiation among organ whorls (calyx/corolla in Cassia; two stamen whorls in Chamaecrista), eccentric initiation on one side of a flower, anther attachment, anther pore structure, and precocious carpel initiation in Senna. The asymmetric corolla and androecium in Chamaecrista arise by precocious organ initiation on one side (left or right). The poricidal anther character can result from differing developmental pathways: lateral slits vs. sealing of lateral sutures; clasping hairs vs. sutural ridges; terminal pores (one or two) vs. none; and clamp layer formation internally that prevents lateral dehiscence. Genera differ in corolla aestivation patterns and in stigma type. Convergence is shown among the three genera, based on intergeneric dissimilarities in early floral ontogeny (floral position in the inflorescence, bracteole presence, position of the first sepal initiated, order of petal initiation, asymmetric initiation, overlap between whorls, anther morphology, and time of carpel initiation) resulting in similarities at anthesis (showy, mostly yellow salverform flowers, heteromorphic stamens, poricidal anther dehiscence, bee pollination, and chambered stigma).

Mutations in the PERIANTHIA gene of Arabidopsis specifically alter floral organ number and initiation pattern.

An open question in developmental biology is how groups of dividing cells can generate specific numbers of segments or organs. We describe the phenotypic effects of mutations in PERIANTHIA, a gene specifically required for floral organ patterning in Arabidopsis thaliana. Most wild-type Arabidopsis flowers have 4 sepals, 4 petals, 6 stamens, and 2 carpels. Flowers of perianthia mutant plants most commonly show a pentamerous pattern of 5 sepals, 5 petals 5 stamens, and 2 carpels. This pattern is characteristic of flowers in a number of plant families, but not in the family Brassicaceae, which includes Arabidopsis. Unlike previously described mutations affecting floral organ number, perianthia does not appear to affect apical or floral meristem sizes, nor is any other aspect of vegetative or floral development severely affected. Floral organs in perianthia arise in a regular, stereotypical pattern similar to that in distantly related species with pentamerous flowers. Genetic analysis shows that PERIANTHIA acts downstream of the floral meristem identity genes and independently of the floral meristem size and floral organ identity genes in establishing floral organ initiation patterns. Thus PERIANTHIA acts in a previously unidentified process required for organ patterning in Arabidopsis flowers.

Inflorescence and Floral Development in Orchidantha maxillarioides (Lowiaceae)

The inflorescence of Orchidantha maxillarioides is similar to a richly branched, polytelic truncate synflorescence. It consists of two to three paracladia borne in the axils of the uppermost phyllomes of the shoot. Each paracladium consists of six bracts. The lowermost bract is the sterile prophyll. The second and third bracts subtend paracladia of the next higher order. The fourth bract subtends a single flower. The fifth and sixth bracts are initiated but do not complete development. Initiation of the paracladia in the axils of bracts 2 and 3 is acropetal, but their differentiation is basipetal. The flowers are trimerous, with a perianth differentiated into sepals and petals. One petal is enlarged to form a labellum. There are five stamens, one inserted opposite each perianth member except the labellum. The ovary is inferior and is closed by a prolongation many times the length of the locular region of the ovary. Floral organ initiation begins with the posterior sepals. The anterior sepal is formed slightly later, at the same time as the initiation of the corolla. The anterior petals arise from primordia that produce only these organs, while the posterior petal (the labellum) arises from a common petal/stamen primordium. The petal/stamen primordium separates into the labellum and the two posterior outer stamens. The anterior outer stamen is initiated slightly later, in the region between the anterior petals. Throughout these stages a floral cup forms below the floral parts. The two inner stamens are initiated simultaneously on the inner surface of this cup, near the insertion of the anterior petals. Three gynoecial primordia are initiated at the top of the floral cup in the region that will form the prolongation. The primordia extend proximally to form the ovary proper and distally to form the style and stigma. The prolongation is formed through intercalary growth. Based on structural and functional similarities in the prolongation and inflorescence, we support placement of the Lowiaceae as the sister group of the Strelitziaceae.