Flower abscission and inflorescence carbohydrates in sensitive and non-sensitive cultivars of grapevine

The Gewurztraminer (GW) and the Pinot noir (PN) cultivars of grapevine differ in their sensitivity to environmental factors that can cause flower abscission, cv. GW being the most sensitive. In order to further define the mechanisms leading to abscission, and owing to the importance of sugars in the achievement of sexual organ ontogenesis, we attempted to correlate the chronology of flower ontogenesis with the variations of carbohydrates in the inflorescence. In the vineyard, under optimal climatic conditions, fruit set of cv. GW and cv. PN was 82% and 65%, respectively. The sugar distribution was different in their inflorescences during the entire duration of flower development. Between stages 15 and 17, flowers of GW and PN reached the crucial meiosis stage. At that time, the inflorescences of cv. GW exhibited higher concentrations of starch and sucrose, whereas those of PN presented higher levels of glucose and fructose. Despite higher starch concentrations in GW inflorescences, starch reserves were present in the ovules and anthers of PN but not in those of GW. These results suggest that the higher content of reserve and transport carbohydrates in the inflorescences of GW favour flower development and fruit set under optimal environmental conditions. Furthermore, since meiosis represents a key step of female development, the different sugar concentrations in the inflorescences of the two cultivars at stages 15 and 17 could be related to the sensitivity to flower abscission under climatic stress. In particular, the presence of starch granules in PN ovules and anthers might explain the higher resistance of this cultivar to flower abscission.     

Distinct regulation in inflorescence carbohydrate metabolism according to grapevine cultivars during floral development

Carbohydrate metabolism is important in plant sexual reproduction because sugar contents are determining factors for both flower initiation and floral organ development. In woody plants, flowering represents the most energy‐consuming step crucial to reproductive success. Nevertheless, in these species, the photosynthesis performed by flowers supplies the carbon required for reproduction. In grapevine (Vitis vinifera), the inflorescence has a specific status because this organ imports carbohydrates at the same time as it exports photoassimilates. In this study, fluctuations in carbohydrate metabolism were monitored by analyzing gas exchanges, photosynthetic electron transport capacity, carbohydrate contents and some activities of carbohydrate metabolism enzymes, in the inflorescences of Pinot noir and Gewurztraminer, two cultivars with a different sensitivity to coulure phenomenon. Our results showed that photosynthetic activity and carbohydrate metabolism are clearly different and differently regulated during the floral development in the two cultivars. Indeed, the regulation of the linear electron flow and the cyclic electron flow is not similar. Moreover, the regulation of PSII activity, with a higher Y(NPQ)/Y(NO) ratio in Gewurztraminer, can be correlated with the higher protection of the photosynthetic chain and consequently with the higher yield under optimal conditions of this cultivar. At least, our results showed a higher photosynthetic activity and a better protection of PSI in Pinot noir during the floral development.     

A molecular genetic perspective of reproductive development in grapevine

The grapevine reproductive cycle has a number of unique features. Inflorescences develop from lateral meristems (anlagen) in latent buds during spring and summer and enter a dormant state at a very immature stage before completing development and producing flowers and berries the following spring. Lateral meristems are unique structures derived from the shoot apical meristem and can either develop into an inflorescence or a tendril. How the grapevine plant controls these processes at the molecular level is not understood, but some progress has been made by isolating and studying the expression of flowering genes in wild-type and mutant grapevine plants. Interestingly, a number of flowering genes are also expressed during berry development. This paper reviews the current understanding of the genetic control of grapevine flowering and the impact of viticulture management treatments and environmental variables on yield. While the availability of the draft genome sequence of grapevine will greatly assist future molecular genetic studies, a number of issues are identified that need to be addressed--particularly rapid methods for confirming gene function and linking genes to biological processes and traits. Understanding the key interactions between environmental factors and genetic mechanisms controlling the induction and development of inflorescences, flowers, and berries is also an important area that requires increased emphasis, especially given the large seasonal fluctuations in yield experienced by the crop and the increasing concern about the effect of climate change on existing wine-producing regions.     

The structure and roles of sterile flowers in Viburnum macrocephalum f. keteleeri (Adoxaceae)

The formation and ecological roles of sterile flowers in flowering plants are interesting issues in floral biology and evolution. Here, we investigated the morphological and anatomical characteristics of both fertile and sterile flowers of Viburnum macrocephalum f. keteleeri, a self-incompatible and insect-pollinated shrub, during different developmental stages of flowers. In addition, pollinator visitation rates and fruit set were determined in intact inflorescences and those with sterile flowers removed. The results indicate that sterile and fertile flowers were developmentally similar during early developmental stages, and that development of the flower types diverged about 15 days before flowering. In addition, pollinator visitation rates, number of pollen grains on stigmas and fruit set were significantly higher in inflorescences with sterile flowers than those without sterile flowers. The results suggest that sterile flowers of this species evolved from fertile flowers under long-term selective pressure, and play a crucial role in enhancing reproductive success through effectively attracting pollinators to the plant and thus enhancing fruit set.     

INFLORESCENCE TYPES AND FRUITING PATTERNS IN HAMLIN AND VALENCIA ORANGES AND MARSH GRAPEFRUIT

The structural, flowering, and fruit-setting patterns of inflorescences of mature Hamlin and Valencia orange and Marsh grapefruit trees were studied for three years. Several development patterns were found, some of which were relatively consistent for the different varieties or years of study. The sequence of anthesis on an inflorescence was: apical flower first, then basal flower, then the subapical flower. Terminals on which the earlier flowers appeared tended to have more inflorescences than those on which flowers appeared later. Inflorescences on which earlier flowers appeared also produced more flowers than those which began flowering later. Inflorescences that began flowering later were more likely to have leaves or have a greater number of leaves than earlier inflorescences. More than half of the inflorescences carried no leaves, and most of these had one flower. No evidence of a relationship between number of flowers and length of the inflorescence was found. Fruit set occurred primarily during the latter part of the flowering period. Many fruit were set on inflorescences without leaves, but on the basis of percent of flowers setting fruit, inflorescences with leaves were more productive. The greatest fruit set occurred in the subapical position on the inflorescence. With growth changes these fruit often appeared to be developing in the apical position. These patterns generally differed little from year to year. Variations may have been due to the differences in the number of flowers produced by the trees. Results were also similar between Hamlin and Valencia oranges. Patterns on Marsh grapefruit resembled those for the oranges but were frequently less consistent.     

Inflorescence of grapevine (Vitis vinifera L.): a high ability to distribute its own assimilates

The distribution of carbon (C) into whole grapevine fruiting cuttings was investigated during flower development to determine the relative contribution of inflorescence and leaf photoassimilates in the total C balance and to investigate their partitioning towards other plant organs. A (13)C labelling procedure was used to label C photoassimilates by leaves and inflorescences in grapevine. Investigations were carried out at various stages of flower/berry development, from separated cluster to fruit set, using grapevine fruiting cuttings with four leaves (Vitis vinifera L. cv. Chardonnay). This is the first study reporting that, during its development, (i) the carbon needs of the inflorescence were met by both leaf and inflorescence photosynthesis, and (ii) the inflorescence amazingly participated significantly to the total C balance of grapevine cuttings by redistributing an important part of its own assimilates to other plant organs. With regard to flowering, 29% of C assimilated by the inflorescence remained in the inflorescence, while partitioning towards the stem reached 42% and, as a lower proportion, 15% in leaves, and 14% in roots.     

Phenology of flowering and starch accumulation in grape (Vitis vinifera L.) cuttings and vines

BACKGROUND AND AIMS: A reliable protocol for flowering and fruiting in cuttings was developed with the aim of (a) studying inflorescence and flower development in grapevine cuttings and field plants, and (b) assisting haploid plant production. METHODS: Inflorescence and flower development was studied in 'Gewurztraminer' (GW) and 'Pinot Noir' (PN) grape vines and cuttings grown in a glasshouse, along with variations in starch in the flowers. As there is a strong relationship between flower development and starch, the starch content of reproductive structures was estimated. KEY RESULTS: Inflorescence and flower development were similar in the vines and cuttings with consistent differences between the two cultivars. Indeed, the ontogenesis of male and female organs is not synchronous in GW and PN, with both female and male meiosis occurring earlier in PN than in GW. Moreover, changes of starch reserves were similar in the two plant types. CONCLUSIONS: Cuttings have a similar reproductive physiology to vines, and can be used to study grape physiology and to develop haploid plants.     

Ecological constraints on fruit initiation by Calyptrogyne ghiesbreghtiana (Arecaceae): FLORAL HERBIVORY,POLLEN AVAILABILITY,AND VISITATION BY POLLINATING BATS

Failure of a flower to initiate fruit can be attributed to an insufficiency of resources, genetic incompatibilities, or ecological constraints on pollination. Flower visitors can increase fruit initiation by pollinating flowers, or decrease fruit initiation by damaging them. If, however, resources limit fruit initiation, variation in visitation may be relatively unimportant. Visitors to Calyptrogyne ghiesbreghtiana, a rain forest understory palm, leave a record of their activity by feeding on floral tissue. I recorded variation in inflorescence visitation, floral display, and fruit initiation during 7 mo over two field seasons. On average, bats visited 50-60% of flowers; katydids and scarab beetles visited and damaged the remaining 40-50%. Four species of bat (Phyllostomidae) captured near inflorescences were found to be carrying C. ghiesbreghtiana pollen. Approximately 18% of flowers visited by bats and not damaged by insects initiated fruit, whereas insect-damaged flowers set a negligible number. Some variation in fruit initiation was explained by nightly variation in the proportion of inflorescences at each site that were potential pollen donors (male-phase inflorescences). Visitors responded to variation in floral display; katydids damaged more flowers on taller inflorescences, and in one season bats visited a greater proportion of flowers on inflorescences with many flowers. However, floral display only explained a small amount of variation in visitation. A similar amount of variation in flower visitation was explained by the degree to which inflorescences were obstructed by surrounding vegetation. I used path analysis to summarize the overall effect of floral display on visitation by both pollinators and florivores and the effect of visitation and site sex ratio on fruit initiation. Because almost all flowers were visited by bats or damaged by katydids, the amount of bat visitation was strongly negatively correlated with katydid damage. In spite of the low fruit set from bat-visited flowers, variation in bat visitation explained 27-37% of the variation in fruit initiation.     

Additive and non-additive effects of simulated leaf and inflorescence damage on survival, growth and reproduction of the perennial herb Arabidopsis lyrata

Herbivores may damage both leaves and reproductive structures, and although such combined damage may affect plant fitness non-additively, this has received little attention. We conducted a 2-year field experiment with a factorial design to examine the effects of simulated leaf (0, 12.5, 25, or 50% of leaf area removed) and inflorescence damage (0 vs. 50% of inflorescences removed) on survival, growth and reproduction in the perennial herb Arabidopsis lyrata. Leaf and inflorescence damage negatively and independently reduced flower, fruit and seed production in the year of damage; leaf damage also reduced rosette size by the end of the first season and flower production in the second year. Leaf damage alone reduced the proportion of flowers forming a fruit and fruit production per plant the second year, but when combined with inflorescence damage no such effect was observed (significant leaf × inflorescence damage interaction). Damage to leaves (sources) caused a greater reduction in future reproduction than did simultaneous damage to leaves and inflorescences (sinks). This demonstrates that a full understanding of the effects of herbivore damage on plant fitness requires that consequences of damage to vegetative and reproductive structures are evaluated over more than 1 year and that non-additive effects are considered.     

The rubber tree (Hevea brasiliensis Muell. Arg.) homologue of the LEAFY/FLORICAULA gene is preferentially expressed in both male and female floral meristems

The rubber tree (Hevea brasiliensis Muell. Arg.) is an important source of natural rubber in tropical regions and, as with many woody species, shows a long juvenile phase. To understand the genetic and molecular mechanisms underlying the reproductive process in rubber trees, H. brasiliensis RRIM600 flower and inflorescence development have been characterized, the rubber tree FLORICAULA/LEAFY (FLO/LFY) orthologue, HbLFY, cloned, and its expression patterns were analysed during vegetative and reproductive development. The rubber tree, similar to other Euphorbiaceae species, produces lateral inflorescences containing male, female, and bisexual flowers. HbLFY is expressed in lateral meristems that give rise to inflorescences and in all flower meristems, consistent with a role in reproductive development. Complementation studies using Arabidopsis lfy mutants indicated that the biological function of LFY might be conserved among Brassicaceae and Euphorbiaceae species.     

FLORAL DISPLAY IN PHYLA INCISA: CONSEQUENCES FOR MALE AND FEMALE REPRODUCTIVE SUCCESS

The inflorescences of Phyla incisa consist of flowers in two phases: younger, nectar-containing flowers that have yellow corolla throats and older, nectar-lacking flowers that have dark purple corolla throats. Observations of pollinator visitation patterns to both natural and manipulated inflorescences were made to determine the role of each flower phase in pollinator attraction. The effect of older-phase flowers on male and female reproductive success was determined by comparing stigmatic pollen loads and estimates of pollen removal from inflorescences having different numbers of these flowers. The pollinators of Phyla selected larger inflorescences more often than expected based upon the size distribution of inflorescences available to them. Both younger- and older-phase flowers contributed to the attraction of pollinators, but the latter were less effective in this function. The presence of older-phase flowers significantly increased the visitation rate to inflorescences and the amount of pollen removed but had little effect on pollen deposition on stigmas. The lack of correspondence between pollen deposition and pollinator-visitation rate was not due to stigma saturation, since stigma loads varied greatly. The data indicate that the deposition of pollen on stigmas in this species is a relatively stochastic process, whereas pollen removal from inflorescences occurs at a much more regular rate. Old-phase flower retention appears to contribute to reproductive success through increased pollen donation when pollinator activity is high and may also increase the probability of seed set when pollinators are rare.     

臭椿花器官分化的初步研究

利用扫描电镜(SEM)和光镜(LM)对臭椿花序及花器官的分化和发育进行了初步研究,表明:1)臭椿花器官分化于当年的4月初,为圆锥花序;2)分化顺序为花萼原基、花冠原基、雄蕊原基和雌蕊原基。5个萼片原基的发生不同步,并且呈螺旋状发生;5个花瓣原基几乎同步发生且其生长要比雄蕊原基缓慢;雄蕊10枚,两轮排列,每轮5个原基的分化基本是同步的;雌蕊5,其分化速度较快;3)在两性花植株中,5个心皮顶端粘合形成柱头和花柱,而在雄株中,5个心皮退化,只有雄蕊原基分化出花药和花丝。本研究着重观察了臭椿中雄花及两性花发育的过程中两性花向单性花的转变。结果表明,臭椿两性花及单性花的形成在花器官的各原基上是一致的(尽管时间上有差异),雌雄蕊原基同时出现在每一个花器官分化过程中,但是,可育性结构部分的形成取决于其原基是否分化成所应有的结构:雄蕊原基分化形成花药与花丝,雌蕊原基分化形成花柱、柱头和子房。臭椿单性花的形成是由于两性花中雌蕊原基的退化所造成,其机理有待于进一步研究。     

Flowering patterns of long-lived Heliconia inflorescences: implications for visiting and resident nectarivores

Summary Flowering patterns of four Heliconia (Heliconiaceae) species in Trinidad, West Indies were examined for their predictability and availability to the nectarivores that rely on Heliconia floral nectar. Principal flower visitors are trapling hermit hummingbirds; inflorescences are inhabited by nectarivorous hummingbird flower mites that move between inflorescences by riding in the hummingbirds' nares. Heliconia inflorescences flower for 40–200 days, providing long-term sources of copious nectar (30–60 l per flower), but each Heliconia flower lasts only a single day. As an inflorescence ages the interval increases between open flowers within a bract; wet-season inflorescences produce open flowers more slowly than dry-season conspecifics.Estimated daily energy expenditures for hermit hummingbirds demonstrate that slow production of short-lived open flowers plus low inflorescence density preclude territorial defense of Heliconia by the hermits. Heliconia flowering patterns are viewed as a means of (i) regulating reproductive investment by the plants through staggered flower production over long periods of time, and (ii) maintaining outcrossing by necessitating a traplining visitation pattern by its hummingbird pollinators. I suggest that Heliconia exhibit a two-tiered pollination system by using hermit hummingbirds primarily for outcrossing and using hummingbird flower mites primarily for self-pollination.     

Nonflowers near the base of extant angiosperms? Spatiotemporal arrangement of organs in reproductive units of Hydatellaceae and its bearing on the origin of the flower

Reproductive units (RUs) of Trithuria, the sole genus of the early-divergent angiosperm family Hydatellaceae, are compared with flowers of their close relatives in Cabombaceae (Nymphaeales). Trithuria RUs combine features of flowers and inflorescences. They differ from typical flowers in possessing an "inside-out" morphology, with carpels surrounding stamens; furthermore, carpels develop centrifugally, in contrast to centripetal or simultaneous development in typical flowers. Trithuria RUs could be interpreted as pseudanthia of two or more cymose partial inflorescences enclosed within an involucre, but the bractlike involucral phyllomes do not subtend partial inflorescences and hence collectively resemble a typical perianth. Teratological forms of T. submersa indicate a tendency to fasciation and demonstrate that the inside-out structure-the primary feature that separates RUs of Hydatellaceae from more orthodox angiosperm flowers-can be at least partially modified, thus producing a morphology that is closer to an orthodox flower. The Trithuria RU could be described as a "nonflower", i.e., a structure that contains typical angiosperm carpels and stamens but does not allow recognition of a typical angiosperm flower. The term nonflower could combine cases of secondary loss of flower identity and cases of a prefloral condition, similar to those that gave rise to the angiosperm flower. Nonhomology among some angiosperm flowers could be due to iterative shifts between nonfloral construction and flower/inflorescence organization of reproductive organs. Potential testing of these hypotheses using evolutionary-developmental genetics is explored using preliminary data from immunolocalization of the floral meristem identity gene LEAFY in T. submersa, which indicated protein expression at different hierarchical levels.     

Vascular development of the grapevine (<Emphasis Type="Italic">Vitis vinifera</Emphasis> L.) inflorescence rachis in response to flower number,plant growth regulators and defoliation

The grapevine inflorescence is a determinate panicle and as buds emerge, shoot, flower and rachis development occur simultaneously. The growth and architecture of the rachis is determined by genetic and environmental factors but here we examined the role of flower and leaf number as well as hormones on its elongation and vascular development. The consequences of rachis morphology and vascular area on berry size and composition were also assessed. One week prior to anthesis, Merlot and Cabernet Sauvignon field vines were exposed to manual flower removal, exogenous plant growth regulators or pre-bloom leaf removal. Manual removal of half the flowers along the vertical axis of the inflorescence resulted in a shorter rachis in both cultivars. Conversely, inflorescences treated with gibberellic acid (GA₃) and the synthetic cytokinin, 6-benzylaminopurine (BAP) resulted in a longer rachis while pre-bloom removal of all leaves on the inflorescence-bearing shoot did not alter rachis length relative to untreated inflorescences. Across the treatments, the cross-sectional areas of the conducting xylem and phloem in the rachis were positively correlated to rachis girth, flower number at anthesis, bunch berry number, bunch berry fresh mass and bunch sugar content at harvest. Conversely, average berry size and sugar content were not linked to rachis vascular area. These data indicate that the morphological and vascular development of the rachis was more responsive to flower number and plant growth regulators than to leaf removal.     

Failure of reproductive assurance in the chasmogamous flowers of Polygala lewtonii (Polygalaceae), an endangered sandhill herb

Hypothetically, a species with both cleistogamous (CL) flowers and delayed selfing chasmogamous (CH) flowers should display high levels of reproductive assurance because, over time, obligate selfing by CL flowers should reduce inbreeding depression and delayed selfing in CH flowers should compensate for the absence of outcross pollen. We used pollinator-exclusion experiments to investigate reproductive assurance in the CH flowers of Polygala lewtonii, an herb with a mixed mating system. We followed CH flowers from bud-break to flower/fruit abscission to quantify fruit initiation and maturation and rates of floral development. We also evaluated the efficacy of the selfing mechanism, conducted pollinator watches to assess the likelihood of pollinator limitation, and performed regression analysis to determine the effect of flower position on fruit production. Pollinator exclusion significantly reduced fruit initiation and maturation. Investigation of floral development demonstrated that the selfing mechanism is largely dysfunctional in CH flowers, indicating the failure of reproductive assurance. Low observed rates of insect visitation appear to contradict high rates of CH fruit production in open-pollinated plants, particularly given the rarity of delayed selfing. In both treatments, flower position significantly affected fruit initiation, suggesting a role for resource limitation in both pollinator-excluded and open-pollinated flowers.     

Reproductive meristem fates in Gerbera

Flowering plants go through several phases between regular stem growth and the actual production of flower parts. The stepwise conversion of vegetative into inflorescence and floral meristems is usually unidirectional, but under certain environmental or genetic conditions, meristems can revert to an earlier developmental identity. Vegetative meristems are typically indeterminate, producing organs continuously, whereas flower meristems are determinate, shutting down their growth after reproductive organs are initiated. Inflorescence meristems can show either pattern. Flower and inflorescence development have been investigated in Gerbera hybrida, an ornamental plant in the sunflower family, Asteraceae. Unlike the common model species used to study flower development, Gerbera inflorescences bear a fixed number of flowers, and the architecture of the flowers differ in that Gerbera ovaries are inferior (borne below the perianth). This architectural difference has been exploited to show that floral meristem determinacy and identity are spatially and genetically distinct in Gerbera, and we have shown that a single SEPALLATA-like MADS domain factor controls both flower and inflorescence meristem fate in the plant. Although these phenomena have not been directly observed in Arabidopsis, the integrative role of the SEPALLATA function in reproductive meristem development may be general for all flowering plants.