Pollinator availability as a determinant of flowering time in ocotillo (Fouquieria splendens)

Summary Ocotillo, a perennial shrub of Sonoran and Chihuahuan Deserts, produces its red tubular flowers in spring. This timing coincides with northward migration of hummingbirds through desert areas. Observations of visitors, pollen collections, and seed set reductions following exclusion of different flower visitors indicate that both hummingbirds and solitary bees pollinate ocotillo in southern Arizona. Seed sets of flowers on marked plants varied considerably within and between years, and this variation was related to the temporal match between flowering and hummingbird migration. This suggests that selection acts on plants to synchronize flowering with periods of pollinator abundance.     

Bumble bee behavior and selection on flower size in the sky pilot,Polemonium viscosum

Summary In alpine Polemonium viscosum, plants having sweet-scented flowers are primarily pollinated by queens of the bumble bee species, Bombus kirbyellus. In this paper we ask whether two aspects of the pollination effectiveness of bumble bees, visitation rate and pollination efficiency, vary significantly with flower size in sweet-flowered P. viscosum.(i) Bumble bees visited plants with large flowers on 80–90% of encounters, but visited those with smaller flowers on only 49% of encounters. (ii) However, the gain in pollination that large-flowered plants obtained via increased visitation was countered in part because bumble bees deposited fewer outcross pollen grains per visit on stigmas of large flowers than on those of small ones. When both visitation rate and pollination efficiency are taken into account, the predicted value of a single bumble bee encounter declines from 1.06 seeds for flowers larger than 18 mm in diameter to 0.55 seeds for flowers smaller than 12 mm in diameter. Our results suggest that bumble bee pollinators of P. viscosum prefer flower morphologies that are poorly suited for precise pollination. Such behavioral complexities are likely to place constraints on the evolution of optimal floral design.     

Decomposition rate of ocotillo (Fouquieria splendens) wood in the desert of southern California and its use in estimating adult survival by life-cycle graph analysis

Dead and downed ocotillos (Fouquieria splendens) are common in the Colorado Desert of southern California and pose a question concerning the life-span of adult plants that have been reported to be long-lived: if ocotillo is long-lived, decay rates must be very slow. Our study focused on decomposition and disintegration of above-ground ocotillo wood over 14 years to obtain a rate of change in dead mass. Disintegration rate was used in a new application of life-cycle graph analysis to calculate persistence time of carcasses, which in turn was used to estimate adult survival rates. We conclude that previous estimates of life spans may be too conservative and half-life of adult shrubs is over 200 years. Our approach to estimating adult survival from decay rates provides an independent test of survival transitions based on live plants for some species and so is an additional tool for developing life cycle models.     

A theory for inflorescence development and flower formation based on morphological and biophysical analysis in Echeveria

Floral development is generally viewed as involving interactions between recently made organs and generative activity on the apical dome; one set of floral organs is thought to induce the next. To investigate such interactions, flowering in Echeveria derenbergii (J. Purpus) was studied at two levels of structure. At the larger, morphological, level the inflorescence apex is shown to have simple cyclic development. Seen from above, it elongates horizontally, then forms a transverse cleft to demarcate a flower primordium in one of two rows. The meristem then elongates at 90° to its previous axis, also horizontally, and demarcates a flower in the other row. Activity on the apical surface correlates well with the nature and activity of adjacent sub-apical organs. For example, the 90° shifts in elongation of the meristem correlate with that tissue's being attached, laterally, to successive large growing bracts whose bases lie at 90°. Also, on the flower primordium, the five sepals arise in a spiral sequence which correlates with one of increasing age, since formation by the cleft, of the edges of the primordium.The second level of study was to test whether the developmental correlations could have a biophysical explanation. By biophysical theory, organs arise where the dome surface is structurally predisposed to bulge. This is a function of the cellulose reinforcement pattern in the surface. Successive patterns of cellulose reinforcement in isolated surface layers from floral organs were determined using polarized light. This was done for the cyclic activity of the inflorescence meristem and the development of the flower. The results indicate that patterns of cellulose reinforcement on the apical dome surface could lead to the production of organs, through local promotion of bulging of the tunica. Subsequent growth of the base of each organ stretches the adjacent dome tissue in a directional fashion. Cytoskeletal responses of these stretched cells lead to new cellulose alignments on the dome which generate the reinforcement pattern for the next round of organs.Abbreviations F floral meristem tissue which will directly produce a flower, starting with sepals - I inflorescence meristem tissue, generally oval in top view and bounded by two bracts, that produces both floral tissue (F) and additional I meristem tissue - I-max the maximal size of I tissue before it bifurcates into F tissue and I tissue (I-min) - I-min the minimal size of I tissue just after it has bifurcated to produce F tissue and I tissue     

Impact of the flower stalk-boring moth Neoschoenobia testacealis (Lepidoptera: Crambidae) and water-level fluctuations on the flower and fruit production of the yellow water lily Nuphar subintegerrima (Nymphaeaceae) in irrigation ponds of western Japan

The flower and fruit production of the yellow water lily Nuphar subintegerrima and the seasonal occurrence of the flower stalk-boring moth Neoschoenobia testacealis were surveyed in two irrigation ponds in western Japan. Water-level fluctuations at each pond were measured. Flowering was observed from late May to September, and fruiting from late May to October. The yellow water lily produced cumulative numbers of 189 and 181 flowers and 121 and 129 fruits in the two ponds’ quadrats. The fruit setting rates at the two ponds were 0.64 and 0.71, and decreased in the summer. The percentage of bored flower stalks was 16 and 5%. Mature larvae bored flower stalks severely during the summer. Whenever the boring larvae attacked the flower stalks, the flower buds and flowers were aborted. There was a significant negative correlation between the fruit setting rate and the boring rate per month (number of bored flower stalks/number of all flower stalks). Whenever abruptly rising water levels (about 10 cm) submerged the flowers, the flowers were aborted too and could not set fruits. The percentages of non-setting fruit due to the rising water levels at the two ponds were 6 and 8%. Overall, our results indicated that flower stalk boring was a more important factor than water-level fluctuation in the sexual reproduction of N. subintegerrima and that N. subintegerrima was well suited to small irrigation ponds.     

Sites of ethylene production in the pollinated and unpollinated senescing carnation (Dianthus caryophyllus) inflorescence

Production of endogenous ethylene from the styles, ovary and petals of pollinated and unpollinated flowers of Dianthus caryophyllus L. was measured. The rate of ethylene production of cut, unpollinated flowers aged in water at 18°C was low until the onset of petal wilting, when a rapid surge of ethylene occurred in all tissues. The flower ethylene production was evolved mostly from the styles and petals. The bases of petals from unpollinated, senescing flowers evolved ethylene faster and sometimes earlier than the upper parts. Treatment of cut flowers with propylene, an ethylene analogue, accelerated wilting of flower petals and promoted endogenous ethylene production in all flower tissues. Pollination of intact flowers also promoted endogenous ethylene production and caused accelerated petal wilting within 2–3 days from pollination. Although the data are consistent with the hypothesis that ethylene forms a link between pollination of the style and petal wilting, in the unpollinated flower the style and petals can evolve a surge of ethylene independently of each other, about the time when the petals irreversibly wilt. The results are discussed in relation to the role of ethylene in flower senescence.     

A procedure for SEM of complex shoot structures applied to the inflorescence of snapdragon (Antirrhinum)

Summary An indirect procedure for the scanning electron microscopy of living complex shoot structures, e.g., an inflorescence tip, employs the polymerization of a dental impression plastic. Application of the plastic to exposed surfaces during prolonged dissection minimizes desiccation. The resulting complex mould is everted so that recesses representing surface detail can be filled with molten epoxy polymer. The mould is then allowed to revert to its original configuration; it is now filled with epoxy. After the epoxy hardens, the resulting cast is sputter-coated and imaged. Intricate structures up to 1 mm in dimension can be imaged with all components, e.g., flowers on an inflorescence, in situ.     

Effects of inflorescence size on visits from pollinators and seed set of Corydalis ambigua (Papaveraceae)

Female reproductive success (seed set) of a spring ephemeral plant, Corydalis ambigua Cham. et schlecht (Papaveraceae) was investigated in relation to inflorescence size and foraging behavior (frequency and duration of visitations) by pollinators (namely, overwintered queens of Bombus hypocrita sapporensis) by detailed daily observations of a natural population. Pollination experiments indicated that C. ambigua is self-incompatible and that seed set was significantly affected by the behavior of the pollinating queens. Plants with larger inflorescences were visited more often than those with fewer flowers. Fecundity also increased with increasing size of inflorescences. Visitation time (duration of foraging) rather than the frequency of visitations (number of visits) was critical for higher fecundity. Seed production was strongly enhanced by a few long visits (of more than 60 s), and seemed to be independent of large numbers of short visits (of less than 60 s). Hence, plants with larger inflorescences, which provide a conspicuous signal to pollinators and offer greater rewards in terms of nectar, received longer visits by B. hypocrita sapporensis queens and those plants exhibited higher fecundity.     

Morphology and anatomy of inflorescence and inflorescence axis in Paepalanthus sect. Diphyomene Ruhland (Eriocaulaceae,Poales) and its taxonomic implications

Paepalanthus sect. Diphyomene has inflorescences arranged in umbels. The underlying bauplan seems however to be more complex and composed of several distinct subunits. Despite appearing superficially very similar, the morphology and anatomy of the inflorescences can supply useful information for the understanding of the phylogeny and taxonomy of the group. Inflorescences of Paepalanthus erectifolius, Paepalanthus flaccidus, Paepalanthus giganteus, and Paepalanthus polycladus were analyzed in regard to branching pattern and anatomy. In P. erectifolius, P. giganteus and P. polycladus the structure is a tribotryum, with terminal dibotryum, and with pherophylls bearing lateral dibotrya. In P. flaccidus, the inflorescence is a pleiobotryum, with terminal subunit, and without pherophylls. Secondary inflorescences may occur in all species without regular pattern. Especially when grown in sites without a pronounced seasonality, the distinction between enrichment zone (part of the same inflorescence) and new inflorescences may be obscured. The main anatomical features supplying diagnostic and phylogenetic information are as follows: (a) in the elongated axis, the thickness of the epidermal cell walls and the cortex size; (b) in the bracts, the quantity of parenchyma cells (c) in the scapes, the shape and the presence of a pith tissue. Therefore, P. sect. Diphyomene can be divided in two groups; group A is represented by P. erectifolius, P. giganteus and P. polycladus, and group B is represented by P. flaccidus. The differentiation is based in both, inflorescence structure and anatomy. Group A presents a life cycle and anatomical features similar to species of Actinocephalus. Molecular trees also point that these two groups are closely related. However, inflorescence morphology and blooming sequence are different. Species of group B present an inflorescence structure and anatomical features shared with many genera and species in Eriocaulaceae. The available molecular and morphology based phylogenies still do not allow a precise allocation of the group in the bulk of basal species of Paepalanthus collocated in P. sect. Variabiles. The characters described and used here supply however important information towards this goal.     

Fruit production and floral traits: correlated evolution in <Emphasis Type="Italic">Govenia</Emphasis> (Orchidaceae)

The most common estimate of reproductive success in orchids is usually fruit set. Factors such as resource limitation and certain floral traits may influence reproductive success in animal-pollinated plants. Correlated evolution of reproductive success vs. seven floral traits (inflorescence length, flower number, flower distribution along inflorescence, dorsal sepal length, lateral sepal length, flower color, and column position) was studied in eight species of Govenia. Taxa represented three lineages in the genus. Independent contrasts were calculated on a phylogeny inferred from chloroplast (trnL-F IGS) DNA sequences, and a correlation test and multiple regression were then performed. Two data sets were evaluated, one including all eight species and another excluding G. utriculata, which is autogamous. The historical analyses showed that there is a correlation between reproductive success and dorsal sepal length, column position, and flower number, these correlations suggest that changes in these floral traits usually accompany evolutionary shifts in reproductive success. Multiple regression tests suggest that changes in reproductive success can be explained by shifts in flower number, inflorescence length, column position and by dorsal sepal length. When phylogeny is taken into account, our analyses showed that evolutionary shifts in these floral traits were correlated with changes in reproductive success. Evolutionary correlation between reproductive success and floral traits might be explained by the natural selection of certain floral phenotypes by pollinators.     

Structure of the inflorescence and flower ofPetalonyx linearis (Loasaceae)

Fossil chloranthoid androecia,Chloranthistemon endressii gen. et spec. nov. are described from the Upper Cretaceous (Upper Santonian or Lower Campanian) of Scania, southern Sweden. They are three-lobed and dorsiventrally flattened with all pollen sacs borne laterally and inclined toward the presumed adaxial surface. The central lobe bears two pairs of pollen sacs, the lateral lobes a single pair each. The morphology, anatomy and valvate dehiscence of the fossil androecia is very similar to that seen in extant species ofChloranthus andSarcandra, but the in situ pollen differs from that of all extantChloranthaceae in being spiraperturate. A single chloranthoid androecium from the Lower Cretaceous (Upper Albian) of Maryland, North America has a more generalized structure thanChloranthistemon endressii. It consists of three stamens that are fused at the base, and each stamen bears two pairs of oppositely positioned pollen sacs. Combined with anatomical information from recentChloranthus the Lower Cretaceous specimen suggests that the androecium in the living genus has arisen by fusion and other modifications of three separate stamens each with a normal complement of four pollen sacs. The structure of both the Upper and Lower Cretaceous androecia suggest that these fossilChloranthaceae were insectpollinated. Macrofossil evidence combined with information from dispersed pollen indicates that theChloranthaceae diversified early in angiosperm fossil history and were an important component of Mid-Cretaceous plant communities.     

水蓼花大小在花序和个体上的变化及其与数量权衡关系研究

探讨一年生植物水蓼花大小在花序上和个体上的变化及花大小与花数量的权衡关系。在54株植物个体上随机选取一花序,在花序的基部、中部和顶部各选取1～2朵花,花大小(生物量)以基部最大(0.851mg),顶部最小(0.715mg),可能是由结构效应引起的。在每个体上随机采集4～20朵花,以其均值表示植物个体的花大小,花大小不随植物个体大小变化而变化。花朵展示和总花数均随个体增大而增加。在控制植物个体大小(地上部分营养器官生物量)后,没有发现花朵展示或花数量与花大小之间的权衡关系,表明个体资源水平的差异可能掩盖了花数量与大小间的权衡关系。     

In vitro propagation of Petasites hybridus (Asteraceae) from leaf and petiole explants and from inflorescence buds

In vitro shoot regeneration from sterile leaf and petiole explants and from in-situ-collected inflorescence buds of Petasites hybridus was achieved by a simple two-step protocol. Murashige and Skoog (MS) nutrient medium was supplemented with 17.6 μm benzyladenine (BA)+0.54 μm naphthaleneacetic acid (NAA) to induce shoots. After 5 weeks of culture, 40% of the petiole and 27% of the leaf explants produced shoots compared to 76% of the inflorescence buds. Single shoots were excised and subcultured on MS medium supplemented with various cytokinins (N⁶-(Δ²-isopentenyl)adenine, BA, kinetin and thidiazuron). A concentration of 8.8 μm kinetin+0.54 μm NAA performed best in terms of shoot multiplication rate, average shoot length and spontaneous root induction. Received: 20 August 1997 / Revision received: 29 December 1997 / Accepted: 5 February 1998     

Activities of auxin polar transport in inflorescence axes of flower mutants ofArabidopsis thaliana: Relevance to flower formation and growth

Relationships between the activity of auxin polar transport and flower formation were studied using several flower mutants ofArabidopsis thaliana. The activity of auxin polar transport in the upper portion of inflorescence axis of wildtype plants ofArabidopsis thaliana was significantly lower than that of the basal part. The activities of auxin polar transport in the upper portion of inflorescence axes ofap1 andclv1 mutants were significantly higher than that of wild-type plant. However, those of other flower mutants tested,ap3-1, ag, pi, Fl-40, Fl-54, Fl-89 andpin-formed, were extremely low as compared with that of wild one. We got some evidence that the reduction of the activity of auxin polar transport is concerned with the growth and development of plants. We could mimic it by the removal of all flowers and pods including mature or immature seeds. Moreover, artificial pollination inap3-1 andpi mutants, in which no seeds are found naturally, resulted in the partial recovery of the activity of auxin polar transport in inflorescence axis. Considering these results in this study together with the fact that inhibitors of auxin polar transport generated almost same disruptions ofpin-formed orpinoid mutants which normally had no flowers in inflorescence axis (Okadaet al. 1991, Uedaet al. 1992, Bennettet al. 1995), the systern of auxin polar transport and its activity in inflorescence axis seems to be essential for the development of flower bud in early stage ofArabidopsis thaliana, and the activity of auxin polar transport is also regulated by the formation of flowers and seeds in inflorescence axis.     

Morphology and development of floral features recognised by pollinators

The diversity of angiosperm flowers is astounding. The conventional explanation for this diversity is that it represents the great variety of ways in which flowers have adapted to attract an even greater diversity of animal pollinators. Many animal behaviourists are therefore interested in how changes in floral morphology affect pollinator behaviour. The establishment of well-characterised model plant species has greatly furthered our understanding of how floral morphology is generated and varied. Many of these model species are pollinated by animals and attract their pollinators through the production of colour, shape, scent, size and rewards. An understanding of the developmental plasticity of floral morphology, and the constraints upon it, should inform research into animal responses to flowers. The use of genetically characterised model species, and the isogenic and near-isogenic lines available in them, will allow dissection of the different components of floral attraction and reward in natural systems. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users. Handling editor: Heikki Hokkanen     

The effect of inflorescence display size and flower position on pollination success in two deceptive and one rewarding orchid

• Inflorescence display size and flower position on the inflorescence play important roles in plant reproduction, in the formation of fruits and are primarily linked to pollinator behaviour. We used three orchids to determine how visitation rates and choice of pollinator depend on number and position of the flowers along the inflorescence.
• We measured reproductive success in (1) natural conditions, (2) hand-pollination experiments and (3) an experimental design, by modifying composition of inflorescences in populations of two deceptive orchids, Orchis anthropophora and O. italica, and one rewarding orchid, Anacamptis coriophora subsp. fragrans.
• There were no differences in natural fruit production in relation to flower position on the inflorescence (i.e. upper versus lower part), suggesting no preference of pollinators for different parts of the inflorescence. Hand-pollination experiments highlighted low pollen limitation in A. coriophora subsp. fragrans but high limitation in O. italica and O. anthropophora. Reproductive success of deceptive orchids in experimental plots decreased significantly when flowers on the upper half of the inflorescence were removed leading to reduced floral display, while reproductive success of the nectariferous species did not differ significantly.
• Our data highlight that in the examined orchids there is no clear relationship between fruit formation and flower position along inflorescences. Thus we can affirm that, for orchids, the entire inflorescence plays a dominant role in insect attraction but the part of the flower spike does not influence the choice of the insect. This implies that all flowers have the same possibility of receiving visits from pollinators, and therefore each flower has the same opportunity to set fruit.

     

Bi-directional inflorescence development inArabidopsis thaliana: Acropetal initiation of flowers and basipetal initiation of paraclades

In this study we investigated Arabidopsis thaliana (L.) Heynh. inflorescence development by characterizing morphological changes at the shoot apex during the transition to flowering. Sixteen-hour photoperiods were used to synchronously induce flowering in vegetative plants grown for 30 d in non-inductive 8-h photoperiods. During the first inductive cycle, the shoot apical meristem ceased producing leaf primordia and began to produce flower primordia. The differentiation of paraclades (axillary flowering shoots), however, did not occur until after the initiation of multiple flower primordia from the shoot apical meristem. Paraclades were produced by the basipetal activation of buds from the axils of leaf primordia which had been initiated prior to photoperiodic induction. Concurrent with the activation of paraclades was the partial suppression of paraclade-associated leaf primordia, which became bract leaves. The suppression of bract-leaf primordia and the abrupt initiation of flower primordia during the first inductive photoperiod is indicative of a single phase change during the transition to flowering in photoperiodically induced Arabidopsis. Morphogenetic changes characteristic of the transition to flowering in plants grown continuously in 16-h photoperiods were qualitatively equivalent to the changes observed in plants which were photoperiodically induced after 30 d. These results suggest that Arabidopsis has only two phases of development, a vegetative phase and a reproductive phase; and that the production of flower primordia, the differentiation of paraclades from the axils of pre-existing leaf primordia and the elongation of internodes all occur during the reproductive phase.