The floral biology of Curcumorpha longiflora (Zingiberaceae): a ginger with two-day flowers

Curcumorpha longiflora is a perennial chasmophyte (rock fissure plant) ginger that usually grows in crevices of calcareous rocks and forms patches on the understory of limestone monsoon rainforests. The pollination ecology of C. longiflora was studied by monitoring phenology and flowering behavior, observing pollinator activity (frequency and behavior of visitors), and the quantity and quality of pollination services. We also investigated the germination of pollen grains and growth of pollen tubes after different pollination treatments to detect its breeding system. Based on the results: (1) for the first time in Zingiberaceae a new protandrous mechanism was found with a two-day flowering to avoid autogamy in this species; (2) under field conditions, all individuals of C. longiflora usually produced only one flower every other day to keep geitonogamy to a minimum; (3) germination of pollen grains and growth rates of pollen tubes under different pollination treatments were the same 4 h later after pollination, suggesting that C. longiflora is completely self-compatible; (4) among the limited visitors, Bombus sp. and Apis florae were effective pollinators, but they were active at different times and at different stages of the flower, probably receiving different rewards.     

Molecular cloning and characterization of NFBP6, a putative floral homeotic gene, from tobacco

 NFBP6 cloned from tobacco is most closely related to petunia FBP6, a putative C-function gene. Also, NFBP6 is expressed specifically in stamens and carpels. Thus, NFBP6 provides a useful tool for molecular characterization of flower development.     

Division of labour within flowers: heteranthery,a floral strategy to reconcile contrasting pollen fates

In many nectarless flowering plants, pollen serves as both the carrier of male gametes and as food for pollinators. This can generate an evolutionary conflict if the use of pollen as food by pollinators reduces the number of gametes available for cross‐fertilization. Heteranthery, the production of two or more stamen types by individual flowers reduces this conflict by allowing different stamens to specialize in ‘pollinating’ and ‘feeding’ functions. We used experimental studies of Solanum rostratum (Solanaceae) and theoretical models to investigate this ‘division of labour’ hypothesis. Flight cage experiments with pollinating bumble bees (Bombus impatiens) demonstrated that although feeding anthers are preferentially manipulated by bees, pollinating anthers export more pollen to other flowers. Evolutionary stability analysis of a model of pollination by pollen consumers indicated that heteranthery evolves when bees consume more pollen than should optimally be exchanged for visitation services, particularly when pollinators adjust their visitation according to the amount of pollen collected.     

Silencing of chaperonin 21, that was differentially expressed in inflorescence of seedless and seeded grapes,promoted seed abortion in tobacco and tomato fruits

Vitis vinifera L. cv. ‘Thompson Seedless’ presents a type of stenospermocarpy in grape where fertilization occurs but seeds abort and fail to develop. To unravel the molecular basis for stenospermocarpy in grapes, subtractive hybridization was carried out in order to isolate differentially regulated genes that participate in the seedlessness machinery. Two ‘Thompson’ lines, a seeded and a seedless, were screened during different flower developmental stages. One of the genes, that was differentially expressed between the seeded and seedless lines, was the chloroplast chaperonin 21 (ch-Cpn21). ch-Cpn21 is a 21-kDa co-chaperonin polypeptide formed by two GroES-like domains fused together in tandem. Silencing of ch-Cpn21 in Nicotiana benthamiana plants resulted in leaf stunting, chlorosis, as well as ovary necrogenesis leading to seed abortion. Moreover, organ-specific silencing of ch-Cpn21 only in Lycopersicum esculentum fruits resulted in the development of seedless tomatoes. These results suggest that ch-Cpn21 may play a role in seed abortion in stenospermocarpic grapes.     

Bumble bees show an induced preference for flowers when primed with caffeinated nectar and a target floral odor

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Interaction between flowers, ants and pollinators: additional evidence for floral repellence against ants

It has been commonly suggested that ants negatively affect plant pollination, particularly in the tropics. We studied ant–flower–pollinator interactions in a lowland rainforest in Borneo. Frequency and duration of pollinator visits were compared between flowers attended by ants and flowers from which ants were excluded. In all four plant species studied, the activity of ants decreased the rate and/or duration of the pollinators’ floral visits. For this and other reasons it is expected that plants repel ants from flowers during anthesis. We tested this prediction for a different set of plant species in which we observed the behaviour of Dolichoderus thoracicus ants when encountering flowers. In eight out of 18 plant species studied, ants showed a significantly higher rejection rate when they encountered flowers than when they encountered controls. Our results are thus consistent with the hypothesis that ants may negatively affect plant fitness by reduced intensity of pollinator visits and that ants are repelled from flowers of many tropical plant species, although this repellence is clearly not ubiquitous.     

Biochemical genetics of resistance to MGBG in tobacco: Mutants that alter SAM decarboxylase or polyamine ratios, and floral morphology

Summary Previously we reported the isolation of Nicotiana tabacum cell lines resistant to methylglyoxal-bis(guanylhydrazone) (MGBG) an inhibitor of the polyamine synthetic enzyme s-adenosylmethionine decarboxylase (SAMde). Here we report that these mutants fall into several distinct classes on the basis of their biochemical characteristics. At least two lines, Mgr12 and Mgr16, have a SAMdc that displays increased resistance to MGBG in in vitro enzyme assays, suggesting that these two alleles have altered the SAMdc enzyme itself, and thus they may serve to identify the SAMdc structural gene. Other cell lines have elevated levels of some of the polyamines or polyamine-conjugates: Mgr11 and Mgr14 have high levels of putrescine and high levels of activity of putrescine synthesizing enzymes; Mgr23 and Mgr3 have elevated spermidine and spermidine-conjugate levels, with Mgr23 also having elevated putrescine levels. Mgr12 and Mgr3 have been analyzed genetically through F₁ crosses with wild-type tobacco, and subsequently by a backcross of an F₁ plant to wild type; however, the total number of seeds obtained in each cross was very small. The results of the genetic analysis are consistent with Mgr12 and Mgr3 being nuclear dominant traits. The floral abnormalities previously reported as associated with these mutations display linkage with the MGBG resistances. At least for Mgr12 we have thus obtained evidence as to the precise nature of the mutation, an altered SAMdc, and demonstrated that this is likely to be genetic cause of the altered flower phenotype.     

Morphological studies on the genusClematis linn. VIII. Floral and inflorescence anatomy inClematis patens with eight-sepaled flowers

The vascular anatomy of inflorescence axes and flowers ofClematis patens have been studied. The species shows a unique behaviour of the vascular bundles in the transition node from vegetative stem to pedicel: stelar bundles increase in number from six to eight as they ascend through the transition node so that the number of vascular bundles coincides with that of sepals. In the pedicel stele the resulting eight bundles are disposed opposite to eight sepals. respectively; each sepal receives its vascular supply from the bundle facing it. Morphological and anatomical evidence suggests that the calyx of eight sepals in this species should be interpreted as having consisted originally of four pairs of opposite organs, rather than as having been derived secondarily through chorisis of sepals from a calyx of four sepals as seen in most other species ofClematis.     

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.     

Combined effects of inflorescence architecture, display size, plant density and empty flowers on bumble bee behaviour: experimental study with artificial inflorescences

Pollen dispersal by pollinators is governed by the extent to which diverse effects on pollinator behaviour act independently or augment or moderate each other. Using artificial inflorescences, we assessed the behavioural responses of bumble bees to inflorescence architecture (raceme, panicle, and umbel), inflorescence size (7 or 13 flowers), inter-inflorescence distance and the proportion of empty flowers per inflorescence. The advantage of large inflorescences in terms of attractiveness was larger for racemes and umbels than for panicles, whereas the effect of inter-inflorescence distance on the number of successive probes was smaller for racemes than for panicles and umbels. The number of flowers probed per visit increased almost proportionally with display size when fewer flowers were empty, whereas the number increased less when many flowers were empty. Our results suggest that display size and the spatial arrangement of flowers and nectar within inflorescences can contribute to efficient pollination by affecting pollinator behaviour interactively.     

Pollen and water limitation in Astragalus scaphoides, a plant that flowers in alternate years

Mast seeding is common in plant populations, but its causes have rarely been tested experimentally. We tested mechanisms of alternate-year flowering and fruit set in an iteroparous, bee-pollinated, herbaceous plant, Astragalus scaphoides, in semi-arid sagebrush steppe. Patterns of reproduction from 1986 to 1999 indicated that spring precipitation was a cue for synchronous flowering, and that increased pollination in high-flowering years was a fitness advantage of synchrony. We tested these patterns by adding supplemental water and pollen to plants in high- and low-flowering sites and years. Supplemental water had no effect on flowering or seed set, so water is not a proximate cue for reproduction, though it could be important over longer (>3 year) time scales. Supplemental pollination increased fruit set in low- but not high-flowering years, indicating that synchronous flowering increases pollination success. Many shorter-term studies also report increased fruit set after pollen supplementation, but not after resource addition. This pattern may reflect the fact that plants can store and reallocate resources, but not pollen, across multiple years. For animal-pollinated herbs such as these, uniting theories about pollination ecology and mast seeding may promote an understanding of the mechanisms that determine patterns of reproduction over time.     

The MADS box gene, FOREVER YOUNG FLOWER, acts as a repressor controlling floral organ senescence and abscission in Arabidopsis

The ectopic expression of a MADS box gene FOREVER YOUNG FLOWER (FYF) caused a significant delay of senescence and a deficiency of abscission in flowers of transgenic Arabidopsis. The defect in floral abscission was found to be due to a deficiency in the timing of cell separation of the abscission zone cells. Down-regulation of INFLORESCENCE DEFICIENT IN ABSCISSION (IDA) may contribute to the delay of the floral abscission in 35S:FYF flowers. FYF was found to be highly expressed in young flowers prior to pollination and was significantly decreased after pollination, a pattern that correlated with its function. Ethylene insensitivity in senescence/abscission and the down-regulation of ETHYLENE RESPONSE DNA-BINDING FACTOR 1 (EDF1) and EDF2, downstream genes in the ethylene response, in 35S:FYF Arabidopsis suggested a role for FYF in regulating senescence/abscission by suppressing the ethylene response. This role was further supported by the fact that 35S:FYF enhanced the delay of flower senescence/abscission in ethylene response 1 (etr1), ethylene-insensitive 2 (ein2) and constitutive triple response 1 (ctr1) mutants, which have defects in upstream genes of the ethylene signaling pathway. The presence of a repressor domain in the C-terminus of FYF and the enhancement of the delay of senescence/abscission in FYF+SRDX (containing a suppression motif) transgenic plants suggested that FYF acts as a repressor. Indeed, in FYF-DR+VP16 transgenic dominant-negative mutant plants, in which FYF was converted to a potent activator by fusion to a VP16-AD motif, the senescence/abscission of the flower organs was significantly promoted, and the expression of BOP2, IDA and EDF1/2 was up-regulated. Our data suggest a role for FYF in controlling floral senescence/abscission by repressing ethylene responses and regulating the expression of BOP2 and IDA in Arabidopsis.     

PROLIFERATING INFLORESCENCE MERISTEM,a MADS-box gene that regulates floral meristem identity in pea

SQUAMOSA and APETALA1 are floral meristem identity genes from snapdragon (Antirrhinum majus) and Arabidopsis, respectively. Here, we characterize the floral meristem identity mutation proliferating inflorescence meristem (pim) from pea (Pisum sativum) and show that it corresponds to a defect in the PEAM4 gene, a homolog of SQUAMOSA and APETALA1. The PEAM4 coding region was deleted in the pim-1 allele, and this deletion cosegregated with the pim-1 mutant phenotype. The pim-2 allele carried a nucleotide substitution at a predicted 5' splice site that resulted in mis-splicing of pim-2 mRNA. PCR products corresponding to unspliced and exon-skipped mRNA species were observed. The pim-1 and pim-2 mutations delayed floral meristem specification and altered floral morphology significantly but had no observable effect on vegetative development. These floral-specific mutant phenotypes and the restriction of PIM gene expression to flowers contrast with other known floral meristem genes in pea that additionally affect vegetative development. The identification of PIM provides an opportunity to compare pathways to flowering in species with different inflorescence architectures.     

Establishment of a CSF-producing cell line from a human gastric carcinoma and characteristics of the CSF produced by that cell line

A human cell line producing colony-stimulating factor has been established in vitro from a human gastric carcinoma. The cell line was transplantable into nude mice which developed a marked neutrophilia. The cell line has been maintained for three years. The cells grew in a monolayered sheet and produced colony-stimulating factors that enhanced the formation of granulocyte and monocyte colonies in vitro with mouse bone marrow cells as the target and granulocyte colonies with human bone marrow cells as the target.     

Desynchronization and re-synchronization of reproduction by Astragalus scaphoides, a plant that flowers in alternate years

Mast seeding, the synchronous seed production by plants at irregular intervals, has been widely studied from the perspective of its fitness benefits, but much less is known about the proximate factors that cause plants to reproduce synchronously. In this article, I follow up on more than two decades of research investigating proximate mechanisms of mast seeding by Astragalus scaphoides, an iteroparous perennial forb. We use long-term monitoring in relation to two environmental manipulations to evaluate the importance of exogenous environmental factors versus endogenous feedbacks for synchrony in this species. Our past research showed that synchrony in this species is explained by the pollen-coupling hypothesis: plants that flower synchronously set seed and deplete stored resources, whereas plants that happen to flower asynchronously are pollen limited, set fewer seeds, and do not deplete resources, and flower again until they are resynchronized. Continued monitoring of two experimental manipulations, water addition, and flower removal, provides additional support for this model, and also reveals subtle effects of water availability on synchrony. Water addition decreased flowering, rather than increasing it as expected based on simple correlations with weather variables, suggesting that precipitation does not synchronize reproduction. Exogenous drivers are generally considered to be the primary synchronizing factors in plant reproduction. Our work in this system suggests that endogenous feedbacks may be more important than has been previously assumed.     

Foraging success of parasitoid wasps on flowers: interplay of insect morphology, floral architecture and searching behavior

Intercropping with flowering herbaceous plants increases parasitoid survivorship, fecundity and retention and pest suppression in agroecosystems. Few studies, however, have examined the compatibility of parasitoid morphology and foraging ability with floral architecture. This study shows that floral architecture influences the selection of floral host resources used to provide nutrients to parasitoids in cropping systems. Parasitoid foraging performance was evaluated using real and artificial flowers which varied in degree of nectar accessibility for two eulophid parasitoids, Edovum puttleri Grissell and Pediobius foveolatus Crawford. Comparisons were made of searching performance on artificial flowers with nectars that were either scented (made from 1:1 honey-water solution) or scentless (made from 1 m sucrose solution) and differences in head widths were compared with corolla apertures. Our results showed a disparity in the ability of E. puttleri and P. foveolatus to gain access to nectar from particular types of floral architectures. E. puttleri fed efficiently only from flowers with exposed nectaries while P. foveolatus foraged efficiently from flowers having either exposed nectaries or nectaries partially obstructed by petals and stamens. Neither wasp species could forage on flowers with cup- or tube-shaped corollas because their heads are wider than the floral apertures. E. puttleri's foraging performance decreased as nectar inaccessibility increased in the artificial flowers, while P. foveolatus' foraging performance was uniform among the different artificial flowers. This indicates that E. puttleri has less propensity to search small openings for nectar than does P. foveolatus. The foraging success of both E. puttleri and P. foveolatus on artificial flowers was lower when 1 M sucrose solution was used as an artificial nectar rather than honey-water solution, indicating that the wasps were stimulated and attracted by the nectar odor. Our systematic evaluation of floral architecture with respect to parasitoid foraging ability has enabled us to predict which types of flowers would serve as suitable floral host plants for parasitoids in the field. That is, only flowers with nectaries that are completely exposed would function as suitable floral host plants for E. puttleri, while P. foveolatus could forage on flowers with either exposed or partially exposed nectaries. Examples of potentially suitable floral hosts suggested from our study include dill (Anethum graveolens L.) and fennel (Foeniculum vulgare L.) for both E. puttleri and P. foveolatus and coriander (Coriandrum sativa L.) for P. foveolatus.     

Identification of a novel hedycaryol synthase gene isolated from Camellia brevistyla flowers and floral scent of Camellia cultivars

Planta - A novel terpene synthase ( Tps ) gene isolated from Camellia brevistyla was identified as hedycaryol synthase, which was shown to be expressed specifically in flowers. Camellia plants are...