Petal Region Research Articles

Structural changes in Nelumbo flower petals during opening and closing.

Nelumbo nucifera is one of several plant species with flowers that typically open in the early morning and close by noon. This movement normally repeats for 3 days, with all petals falling off on day 4. However, detailed observations of flower movement in Nelumbo species are limited. Movement of flowers on plants in growth chambers were observed using time-lapse photography. Petals were examined with scanning electron microscopy or fixed and sectioned for light microscopy to determine whether changes in cell size contribute to petal elongation during flowering. This study is the first to microscopically observe petal cells of a Nelumbo species during flowering in controlled conditions. Petals elongated during the 4-day flowering period. Inner petals were more elongated than the outer petals. Among the basal, central, and tip regions of a single petal, the cells in the basal region enlarged the most. Outer and inner epidermal cells in the basal region of the inner petals, on both adaxial and abaxial sides, gradually enlarged during the flowering period through cycles of repeated increases as the petals opened and decreases as they closed. Petals opened and closed repeatedly as they elongated, primarily in their basal region. Cells in the basal region of the petal on the adaxial and abaxial sides periodically fluctuated in size, increased during flower opening and decreased during closing, but differences in these patterns were observed between the two sides, suggesting that one of the driving forces behind the opening and closing of flowers of Nelumbo species is the increase and decrease in the size of the petal cells.

Hibiscus bullseyes reveal mechanisms controlling petal pattern proportions that influence plant-pollinator interactions.

Colorful flower patterns are key signals to attract pollinators. To produce such motifs, plants specify boundaries dividing petals into subdomains where cells develop distinctive pigmentations, shapes, and textures. While some transcription factors and biosynthetic pathways behind these characteristics are well studied, the upstream processes restricting their activities to specific petal regions remain enigmatic. Here, we unveil that the petal surface of Hibiscus trionum, an emerging model featuring a bullseye on its corolla, is prepatterned as the bullseye boundary position is specified long before it becomes visible. Using a computational model, we explore how pattern proportions are maintained while petals experience a 100-fold size increase. Exploiting transgenic lines and natural variants, we show that plants can regulate boundary position during the prepatterning phase or modulate growth on either side of this boundary later in development to vary bullseye proportions. Such modifications are functionally relevant, as buff-tailed bumblebees can reliably identify food sources based on bullseye size and prefer certain pattern proportions.

Complex petal spot formation in the Beetle Daisy (Gorteria diffusa) relies on spot-specific accumulation of malonylated anthocyanin regulated by paralogous GdMYBSG6 transcription factors.

Gorteria diffusa has elaborate petal spots that attract pollinators through sexual deception, but how G. diffusa controls spot development is largely unknown. Here, we investigate how pigmentation is regulated during spot formation. We determined the anthocyanin composition of G. diffusa petals and combined gene expression analysis with protein interaction assays to characterise R2R3-MYBs that likely regulate pigment production in G. diffusa petal spots. We found that cyanidin 3-glucoside pigments G. diffusa ray floret petals. Unlike other petal regions, spots contain a high proportion of malonylated anthocyanin. We identified three subgroup 6 R2R3-MYB transcription factors (GdMYBSG6-1,2,3) that likely activate the production of spot pigmentation. These genes are upregulated in developing spots and induce ectopic anthocyanin production upon heterologous expression in tobacco. Interaction assays suggest that these transcription factors regulate genes encoding three anthocyanin synthesis enzymes. We demonstrate that the elaboration of complex spots in G. diffusa begins with the accumulation of malonylated pigments at the base of ray floret petals, positively regulated by three paralogous R2R3-MYB transcription factors. Our results indicate that the functional diversification of these GdMYBSG6s involved changes in the spatial control of their transcription, and modification of the duration of GdMYBSG6 gene expression contributes towards floral variation within the species.

Petal Region Analysis of Improved Petal Ant Routing for Mobile Ad Hoc Networks

Petal Region Analysis of Improved Petal Ant Routing for Mobile Ad Hoc Networks

Transcriptome Analysis Reveals the Molecular Mechanisms Associated with Flower Color Formation in Camellia japonica ‘Joy Kendrick’

Camellia japonica is a woody flower with high ornamental and economic value used for landscaping and as a pot plant. Floral colors are among the most important ornamental traits of flower plants, particularly multicolored flowers. The C. japonica cultivar ‘Joy Kendrick’ has multicolored flowers; the corolla is pink with darker red stripes, but the molecular mechanism underlying this trait is unknown. Here, pigment analysis showed that there are more anthocyanins accumulate in red petal regions than in pink areas, which may be key to formation of red stripes. Furthermore, transcriptome analysis revealed that anthocyanin biosynthesis, modification, and transporter genes are highly expressed in red stripes, consistent with the observed anthocyanin accumulation. In addition, many plant hormone signal transduction genes, particularly auxin, may contribute to the regulation of red stripe formation. This study provides broad insights into pigment accumulation and the regulatory mechanisms underlying floral color formation in C. japonica, and lays a foundation for breeding new C. japonica varieties.

Arabidopsis transcription factor TCP4 represses chlorophyll biosynthesis to prevent petal greening

Green petals pose a challenge for pollinators to distinguish flowers from leaves, but they are valuable as a specialty flower trait. However, little is understood about the molecular mechanisms that underlie the development of green petals. Here, we report that CINCINNATA (CIN)-like TEOSINTE BRANCHED 1/CYCLOIDEA/PCF (TCP) proteins play key roles in the control of petal color. The septuple tcp2/3/4/5/10/13/17 mutant produced flowers with green petals due to chlorophyll accumulation. Expression of TCP4 complemented the petal phenotype of tcp2/3/4/5/10/13/17. We found that chloroplasts were converted into leucoplasts in the distal parts of wild-type petals but not in the proximal parts during flower development, whereas plastid conversion was compromised in the distal parts of tcp2/3/4/5/10/13/17 petals. TCP4 and most CIN-like TCPs were predominantly expressed in distal petal regions, consistent with the green–white pattern in wild-type petals and the petal greening observed in the distal parts of tcp2/3/4/5/10/13/17 petals. RNA-sequencing data revealed that most chlorophyll biosynthesis genes were downregulated in the white distal parts of wild-type petals, but these genes had elevated expression in the distal green parts of tcp2/3/4/5/10/13/17 petals and the green proximal parts of wild-type petals. We revealed that TCP4 repressed chlorophyll biosynthesis by directly binding to the promoters of PROTOCHLOROPHYLLIDE REDUCTASE (PORB), DIVINYL REDUCTASE (DVR), and SUPPRESSOR OF OVEREXPRESSION OF CO 1 (SOC1), which are known to promote petal greening. We found that the conversion of chloroplasts to leucoplasts and the green coloration in the proximal parts of petals appeared to be conserved among plant species. Our findings uncover a major molecular mechanism that underpins the formation of petal color patterns and provide a foundation for the breeding of plants with green flowers.

Structure of the bacterial flagellar hook cap provides insights into a hook assembly mechanism

Assembly of bacterial flagellar hook requires FlgD, a protein known to form the hook cap. Symmetry mismatch between the hook and the hook cap is believed to drive efficient assembly of the hook in a way similar to the filament cap helping filament assembly. However, the hook cap dependent mechanism of hook assembly has remained poorly understood. Here, we report the crystal structure of the hook cap composed of five subunits of FlgD from Salmonella enterica at 3.3 Å resolution. The pentameric structure of the hook cap is divided into two parts: a stalk region composed of five N-terminal domains; and a petal region containing five C-terminal domains. Biochemical and genetic analyses show that the N-terminal domains of the hook cap is essential for the hook-capping function, and the structure now clearly reveals why. A plausible hook assembly mechanism promoted by the hook cap is proposed based on the structure.

Implications of region-specific gene expression for development of the partially fused petunia corolla.

Angiosperm petal fusion (sympetaly) has evolved multiple times independently and is associated with increased specificity between plants and their pollinators. To uncover developmental genetic changes that might have led to the evolution of sympetaly in the asterid core eudicot genus Petunia (Solanaceae), we carried out global and fine-scale gene expression analyses in different regions of the corolla. We found that, despite several similarities with the choripetalous model species Arabidopsis thaliana in the proximal-distal transcriptome, the Petunia axillaris fused and proximal corolla tube expresses several genes that in A. thaliana are associated with the distal petal region. This difference aligns with variation in petal shape and fusion across ontogeny of the two species. Moreover, differential gene expression between the unfused lobes and fused tube of P.axillaris petals revealed three strong candidate genes for sympetaly based on functional annotation in organ boundary specification. Partial silencing of one of these, the HANABA TARANU (HAN)-like gene PhGATA19, resulted in reduced fusion of Petunia hybrida petals, with silencing of both PhGATA19 and its close paralog causing premature plant senescence. Finally, detailed expression analyses for the previously characterized organ boundary gene candidate NO APICAL MERISTEM (NAM) supports the hypothesis that it establishes boundaries between most P.axillaris floral organs, with the exception of boundaries between petals.

Transcriptome analysis reveals the regulation of brassinosteroids on petal growth in Gerbera hybrida.

Gerbera hybrida is a cut-flower crop of global importance, and an understanding of the mechanisms underlying petal development is vital for the continued commercial development of this plant species. Brassinosteroids (BRs), a class of phytohormones, are known to play a major role in cell expansion, but their effect on petal growth in G. hybrida is largely unexplored. In this study, we found that the brassinolide (BL), the most active BR, promotes petal growth by lengthening cells in the middle and basal regions of petals, and that this effect on petal growth was greater than that of gibberellin (GA). The RNA-seq (high-throughput cDNA sequencing) technique was employed to investigate the regulatory mechanisms by which BRs control petal growth. A global transcriptome analysis of the response to BRs in petals was conducted and target genes regulated by BR were identified. These differentially expressed genes (DEGs) include various transcription factors (TFs) that were activated during the early stage (0.5 h) of BL treatment, as well as cell wall proteins whose expression was regulated at a late stage (10 h). BR-responsive DEGs are involved in multiple plant hormone signal pathways, hormone biosynthesis and biotic and abiotic stress responses, showing that the regulation of petal growth by BRs is a complex network of processes. Thus, our study provides new insights at the transcriptional level into the molecular mechanisms of BR regulation of petal growth in G. hybrida.

Genetic architecture, biochemical underpinnings and ecological impact of floral UV patterning.

Floral attraction traits can significantly affect pollinator visitation patterns, but adaptive evolution of these traits may be constrained by correlations with other traits. In some cases, molecular pathways contributing to floral attraction are well characterized, offering the opportunity to explore loci potentially underlying variation among individuals. Here, we quantify the range of variation in floral UV patterning (i.e. UV 'bulls-eye nectar guides) among crop and wild accessions of Brassica rapa. We then use experimental crosses to examine the genetic architecture, candidate loci and biochemical underpinnings of this patterning as well as phenotypic manipulations to test the ecological impact. We find qualitative variation in UV patterning between wild (commonly lacking UV patterns) and crop (commonly exhibiting UV patterns) accessions. Similar to the majority of crops, recombinant inbred lines (RILs) derived from an oilseed crop × WI fast-plant® cross exhibit UV patterns, the size of which varies extensively among genotypes. In RILs, we further observe strong statistical-genetic and QTL correlations within petal morphological traits and within measurements of petal UV patterning; however, correlations between morphology and UV patterning are weak or nonsignificant, suggesting that UV patterning is regulated and may evolve independently of overall petal size. HPLC analyses reveal a high concentration of sinapoyl glucose in UV-absorbing petal regions, which, in concert with physical locations of UV-trait QTLs, suggest a regulatory and structural gene as candidates underlying observed quantitative variation. Finally, insects prefer flowers with UV bulls-eye patterns over those that lack patterns, validating the importance of UV patterning in pollen-limited populations of B. rapa.

Vasculature pattern of the petals of the species Kallstroemia pubescens, which indicates the evolutionary status of this taxa

This paper deals with the details variation of petal venation in Kallstroemia pubescens of the family Zygophyllaceae. In the present study, from a pool of 100 petals, 40 petals were selected randomly to observe the variation of venation patterns of petals in different flowers of same species collected from different population of the same ecological area. All the 40 petals examined, as per Foster’s classification, shows various type of anastomoses. Detailed study reveals that after the entry of a single vain in a petal, it divided into two costals and one median. Actually anastomosis takes place between the sister’s branches of the costal or between a branch of the costal and a branch of the medion. Present observation on the anastomosis of the petal venation of the K. pubescens is clearly fits with the study of Foster (1968). At the basal region of petal, Type-I(35.2%) and Type-II(35.2%) are more prevalent, than Type-III (5.8%), Type-VI (5.8%) and Type-V (17.5%).At the middle region of petal, Type-II (47.56%) is more prevalent than Type-I (17.07%), Type-V (13.41%), Type-III (13.41%), Type-IV (4.8%) and Type-VI (3.65%). At the distal region, Type-II (44.7%) is more prevalent than Type-III (33.33%), Type-I (10.41%), Type-V (8.33 %), Type-VI (2.08 %) and Type-IV (1.04 %).

Transmission Fourier transform infrared microspectroscopy allows simultaneous assessment of cutin and cell‐wall polysaccharides of Arabidopsis petals

A procedure for the simultaneous analysis of cell-wall polysaccharides, amides and aliphatic polyesters by transmission Fourier transform infrared microspectroscopy (FTIR) has been established for Arabidopsis petals. The combination of FTIR imaging with spectra derivatization revealed that petals, in contrast to other organs, have a characteristic chemical zoning with high amount of aliphatic compounds and esters in the lamina and of polysaccharides in the stalk of the petal. The hinge region of petals was particular rich in amides as well as in vibrations potentially associated with hemicellulose. In addition, a number of other distribution patterns have been identified. Analyses of mutants in cutin deposition confirmed that vibrations of aliphatic compounds and esters present in the lamina were largely associated with the cuticular polyester. Calculation of spectrotypes, including the standard deviation of intensities, allowed detailed comparison of the spectral features of various mutants. The spectrotypes not only revealed differences in the amount of polyesters in cutin mutants, but also changes in other compound classes. For example, in addition to the expected strong deficiencies in polyester content, the long-chain acyl CoA synthase 2 mutant showed increased intensities of vibrations in a wavelength range that is typical for polysaccharides. Identical spectral features were observed in quasimodo2, a cell-wall mutant of Arabidopsis with a defect in pectin formation that exhibits increased cellulose synthase activity. FTIR thus proved to be a convenient method for the identification and characterization of mutants affected in the deposition of cutin in petals.

Mass spectrometric imaging as a high-spatial resolution tool for functional genomics: Tissue-specific gene expression of TT7 inferred from heterogeneous distribution of metabolites in Arabidopsis flowers

Laser desorption/ionization (LDI) mass spectrometry imaging (MSI) was used to acquire chemical images of flavonoid metabolites on the surface of wild-type and mutant (tt7) Arabidopsis thaliana flowers. Flavonoids were localized to the petals and carpels of flowers, with tissue heterogeneity in the petals. Specifically, kaempferol and/or its glycosides were abundant in the distal region of petals and quercetin and its downstream flavonoids were highly enriched in the more proximal region of petals. As a result of a mutation in the TT7 gene which blocks the conversion of dihydrokaempferol to dihydroquercetin, the downstream metabolites, quercetin, isohamnetin, and their glycosides, were not observed in the mutant flowers. Instead, the metabolites in an alternative pathway, kaempferol and/or its glycosides, were as highly abundant on the proximal region of the petals as in the distal region. In addition, the combined flavonoid amounts on the proximal region of petals in the wild-type are almost equivalent to the amounts of kaempferol and/or its glycosides in the mutant. This strongly suggests that the expression of the TT7 gene is localized on the proximal part of the petal while the other genes in the upper stream pathway are evenly expressed throughout the petal. Most importantly, this work demonstrates MSI of metabolites can be utilized for the localization of gene expression.

Three-dimensional formation of corolla shapes in relation to the developmental distortion of petals in Eustoma grandiflorum

Associations between petal growth patterns and the formation of three-dimensional corolla shapes were investigated using five cultivars of Eustoma grandiflorum with different corolla shapes. The cultivars showing outward curvature in the distal regions of petals exhibited rapid distal petal widening during the later stages of flower opening, whereas the cup-shaped cultivars exhibited rapid widening in the basal to middle regions of petals during earlier stages of flower development. In a funnel-shaped cultivar showing no apparent curvature, no regional differences in petal expansion were found. Distal petal widening was not due to cell widening but was the result of the rapid expansion both in width and length of petal cells, whereas basal expansion was partly due to cell widening. The data obtained suggested that non-uniform cell expansion within a petal resulted in petal distortion, and this played a central role in corolla curvature of E. grandiflorum.

The mutants compacta ähnlich, Nitida and Grandiflora define developmental compartments and a compensation mechanism in floral development in Antirrhinum majus

In order to improve our understanding of floral size control we characterised three mutants of Antirrhinum majus with different macroscopic floral phenotypes. The recessive mutant compacta ähnlich has smaller flowers affected mainly in petal lobe expansion, the dominant mutant Grandiflora has overall larger organs, whilst the semidominant mutation Nitida exhibits smaller flowers in a dose-dependent manner. We developed a cell map in order to establish the cellular phenotypes of the mutants. Changes in organ size were both organ- and region-specific. Nitida and compacta ähnlich affected cell expansion in proximal and distal petal regions, respectively, suggesting differential regulation between petal lobe regions. Although petal size was smaller in compacta ähnlich than in wild type, conical cells were significantly bigger, suggesting a compensation mechanism involved in petal development. Grandiflora had larger cells in petals and increased cell division in stamens and styles, suggesting a relationship between genes controlling organ size and organ identity. The level of ploidy in petals of Grandiflora and coan was found to be equivalent to wild type petals and leaves, ruling out an excess of growth via endoreduplication. We discuss our results in terms of current models about control of lateral organ size.

Identification and preliminary analysis of a new PCP promoter from Brassica rapa ssp. chinensis

The promoter of Brassica campestris Male Fertile 5 (BcMF5), a pollen coat protein member, class A (PCP-A) gene family, was isolated from Brassica rapa L. ssp. chinensis Makino (Chinese cabbage-pak-choi) by Thermal Asymmetric Interlaced Polymerase Chain Reaction (TAIL-PCR). Sequence analysis suggested that the 605-bp promoter of BcMF5 appears to be a pollen promoter. In an attempt to confirm the promoter activity of BcMF5 promoter, -609 to +3 bp and -377 to +3 bp fragments of the upstream sequence of BcMF5 were inserted at the site upstream of the coding region of the uidA gene in the sense orientation to construct two deletion expression vectors. Transient expression analysis in onion epidermal cells by particle bombardment showed that both -609 to +3 bp and -377 to +3 bp fragments of BcMF5 promoter were capable of driving beta-glucuronidase gene expression. Furthermore, by Agrobacterium-mediated genetic transformation method, Arabidopsis transgenic Kan(R) plants were obtained. GUS assay analysis revealed that the promoter of BcMF5 induced gene expression at the early stage of anther development and drove high levels of GUS expression in anther walls, upper regions of petals, pollen, and pollen tubes in the middle and late stage of anther development, but did not drive any expression in sepals and pistils.

The F-box-containing protein UFO and AGAMOUS participate in antagonistic pathways governing early petal development in Arabidopsis.

The UNUSUAL FLORAL ORGANS (UFO) gene is required for multiple processes in the developing Arabidopsis flower, including the proper patterning and identity of both petals and stamens. The gene encodes an F-box-containing protein, UFO, which interacts physically and genetically with the Skp1 homolog, ASK1. In this report, we describe four ufo alleles characterized by the absence of petals, which uncover another role for UFO in promoting second whorl development. This UFO-dependent pathway is required regardless of the second whorl organ to be formed, arguing that it affects a basic process acting in parallel with those establishing organ identity. However, the pathway is dispensable in the absence of AGAMOUS (AG), a known inhibitor of petal development. In situ hybridization results argue that AG is not transcribed in the petal region, suggesting that it acts non-cell-autonomously to inhibit second whorl development in ufo mutants. These results are combined into a genetic model explaining early second whorl initiation/proliferation, in which UFO functions to inhibit an AG-dependent activity.

A floral boost.

A floral boost.

Spatial fragrance patterns within the flowers ofRanunculus acris (Ranunculaceae)

Floral scents emitted from different flower parts ofRanunculus acris were investigated by trapping headspace volatiles onto Porapak Q followed by solvent desorption and GC-MS analysis. Isoprenoids, strongly dominated bytrans-β-ocimene, constituted the principal class of volatiles in all flower parts except pollen; sesquiterpenes were especially diverse. Odors collected separately from petals, stamens, and sepals + gynoecium comprised the same volatiles, but these were present in disparate proportions among the flower parts, thereby creating subtle contrasts within the flower. The main sources of volatiles were the petals and stamens, which made equal contributions to the whole-flower fragrance. Emissions from the petals differed quantitatively between the apical and basal petal regions, thereby paralleling optical nectar-guide patterns. Pollen odor was markedly unlike that of other flower parts, with only few volatiles, a high representation of 5-methylene-2(5H)-furanone (protoanemonin), and no detectabletrans-β-ocimene. The distinctiveness of the pollen's volatile profile suggests that it may serve a signalling role to pollen-feeding insects.

Structure and ontogeny of stomata on the vegetative and floral organs in nine species ofVerbena

The structure and ontogeny of stomata on the vegetative and floral organs in nine species ofVerbena are described. The data on stomatal index, density and size and epidermal cell density and size are also tabulated. Nine types of stomata occur side by side on the same surface in different combinations. The stomatal ontogeny is mesogenous, mesoperigenous and perigenous. Stomata with common subsidiary cells, cytoplasmic connections, cuticular striations, unequal guard cells, one guard cell and contiguous stomata are but sparingly observed. An increase in the number of subsidiary cells takes place by wall formation or the neighbouring perigenes assuming their shapes. Epidermal cells with inward projections are observed in the limb region of petals in all species. The taxonomic significance of stomata is also indicated.