POLLINATION IN FLORAL SCENT MORPHS OF POLEMONIUM VISCOSUM: A MECHANISM FOR DISRUPTIVE SELECTION ON FLOWER SIZE

Plants of Polemonium viscosum have flowers that are either sweet or skunky in scent. The two morphs are preferentially pollinated by insects of strongly contrasting body size: bumblebee queens specialize on sweet flowers, flies on skunky ones. In this study 13 characters were examined in plant specimens from five populations to identify major components of intraspecific variation in flower and inflorescence morphology and test their correlation with floral scent. Factor analysis identified four major axes of morphological variation. The first explained 22% of the variance among specimens and correlated strongly with four flower size characters: sepal length, corolla tube length, corolla lobe width, and corolla lobe length. Floral scent morphs differed significantly in the multivariate representation of flower size defined by these characters. Sweet flowers had wider corolla lobes, longer corolla tubes, and longer sepals than skunky ones. Corolla lobe width accounted for the greatest amount of intermorph divergence. Divergence in flower size between morphs was maintained in mixed populations at four locations in alpine Colorado, with corollas of sweet flowers significantly broader or more flared than those of skunky flowers. Patterns of pollen receipt suggest that this difference is adaptive. In the sweet morph, pollination intensity and purity increased significantly with corolla flare. Conversely, in the skunky morph, corolla flare had little influence on pollination intensity and had a strong negative effect on purity. These findings suggest that selection for effective pollination should favor intraspecific divergence in flower size in Polemonium viscosum.     

APICAL SIZE IN THE DETERMINATION OF COROLLA LOBE NUMBER IN LINANTHUS ANDROSAECUS SSP. ANDROSAECUS

A hypothesis in the literature suggests that apical size is one of the critical determinants of the number of organs a flower will form. Genetically selected and control lines of Linanthus androsaecus ssp. androsaecus (Polemoniaceae) were used to test this hypothesis. Plants in the selection up (SU) line of this species produce flowers with greater than the normal number of corolla lobes, and plants in the selection down (SD) line produce less than the normal number of corolla lobes. Apices from lateral branches collected from plants just prior to flowering differed in size among these two lines and controls. Plants in the SU line showed an increase in apical diameter in median section while plants in the SD line showed a decrease in apical diameter in comparison to controls. SU plants may be flowering later, and SD plants earlier, than controls. SU and control apices from the main stem, collected from plants in a late stage of vegetative growth, also differed in apical diameter. Increase in the number of apical cells is associated with increased size in SU apices in comparison to controls. Apical size differences associated with petal number appear to become established during vegetative growth by the time the first two pairs of foliage leaves are formed. The selected and control lines of L. androsaecus offer support for the hypothesis that apical size is an important factor in the determination of petal number.     

FLORAL ORGANOGENESIS IN FIVE GENERA OF THE MARANTACEAE AND IN CANNA (CANNACEAE)

The paired flowers of all species of the Marantaceae studied, except Monotagma plurispicatum, are produced through the division of an apical meristem with a tunica-corpus structure. The solitary flowers of M. plurispicatum develop from a similar meristem which does not bifurcate. The paired flowers of Canna indica are produced in the axil of a florescence bract through the formation of a bract and an axillary flower on the side of the primordium which gives rise to the largest flower of the pair. The sequence of organ initiation for both families is: calyx, corolla and inner androecial whorl, outer androecial whorl, gynoecium. The sequence of sepal formation is opposite in the two families. In the Cannaceae it leads directly into the spiral created by the formation of the other organs, while in the Marantaceae the sequence of sepal formation follows a spiral opposite to that of the other floral organs. The members of the corolla and inner androecial whorl separate from common primordia. In general these common primordia separate into a petal and an inner androecial member through the initiation of two growth centers, at the same level, in the dorsal and ventral flanks of the primordium. In Ischnosiphon elegans and Pleiostachya pruinosa the stamen is initiated at a lower position than the petal in the ventral flank of the common primordium. A similar pattern of initiation is described for the callose staminode in Marantochloa purpurea and Canna indica. This pattern is interpreted as a variation on the more generalized pattern of inner androecial formation found in the other genera.     

Trehalose induces the ADP-glucose pyrophosphorylase gene, ApL3, and starch synthesis in Arabidopsis

In Arabidopsis, genes encoding functional enzymes for the synthesis and degradation of trehalose have been detected recently. In this study we analyzed how trehalose affects the metabolism and development of Arabidopsis seedlings. Exogenously applied trehalose (25 mM) strongly reduced the elongation of the roots and, concomitantly, induced a strong accumulation of starch in the shoots, whereas the contents of soluble sugars were not increased. When Arabidopsis seedlings were grown on trehalose plus sucrose (Suc), root elongation was restored, but starch still accumulated to a much larger extent than during growth on Suc alone. The accumulation of starch in the shoots of trehalose-treated seedlings was accompanied by an increased activity of ADP-glucose pyrophosphorylase and an induction of the expression of the ADP-glucose pyrophosphorylase gene, ApL3. Even in the presence of 50 mM Suc, which itself also slightly induced ApL3, trehalose (5 mM) led to a further increase in ApL3 expression. These results suggest that trehalose interferes with carbon allocation to the sink tissues by inducing starch synthesis in the source tissues. Furthermore, trehalose induced the expression of the beta-amylase gene, AT-beta-Amy, in combination with Suc but not when trehalose was supplied alone, indicating that trehalose can modulate sugar-mediated gene expression.     

Approaches to influence starch quantity and starch quality in transgenic plants

Starch plays a major role as a transitory and long-term storage compound in higher plants, and therefore is of prime importance for plant growth and development. Additionally, starch serves as a widely used material for a variety of industrial uses. The formation of starch can arbitrarily be divided into three types of event: (I) those leading to the supply of glucose-1-phosphate in the plastids; (II) the conversion of glucose-1-phosphate to ADP-glucose catalysed by the enzyme ADP-glucose pyrophosphorylase; and (III) the enzymatic reactions converting ADP-glucose to long-chain glucans (amylopectin, amylose). In recent years, numerous cDNA and genomic sequences encoding enzymes involved in starch metabolism have been identified. Some of these have been used to down-regulate enzyme activities via the antisense RNA technique. Additionally, bacterial genes have been ectopically expressed in transgenic plants in order to increase corresponding enzyme activities. By modulating the activity of ADP-glucose pyrophosphorylase in plastids, it was possible to decrease and increase, respectively, the starch content in source and sink organs of transgenic plants. In addition, down-regulation of granule-bound starch synthase (isoform I) resulted in the production of starch that was almost completely free of amylose. Further experiments aimed to modulate starch structure are currently underway and will briefly be discussed.     

Spatial and temporal distribution of mineral nutrients and sugars throughout the lifespan of <Emphasis Type="Italic">Hibiscus rosa-sinensis</Emphasis> L. flower

Although the physiological and molecular mechanisms of flower development and senescence have been extensively investigated, a whole-flower partitioning study of mineral concentrations has not been carried out. In this work, the distribution of sucrose, total reducing sugars, dry and fresh weight and macro and micronutrients were analysed in Hibiscus rosa-sinensis L. petals, stylestigma including stamens and ovary at different developmental stages (bud, open and senescent flowers). Total reducing sugars showed the highest value in petals of bud flowers, then fell during the later stages of flower development whereas sucrose showed the highest value in petals of senescent flowers. In petals, nitrogen and phosphorus content increased during flower opening, then nitrogen level decreased in senescent flowers. The calcium, phosphorus and boron concentrations were highest in ovary tissues whatever the developmental stage. Overall, the data presented suggests that the high level of total reducing sugars prior the onset of flower opening contributes to support petal cells expansion, while the high amount of sucrose at the time of petal wilting may be viewed as a result of senescence. Furthermore, this study discusses how the accumulation of particular mineral nutrients can be considered in a tissue specific manner for the activation of processes directly connected with reproduction.     

Developmental anatomy of foliage leaves,bracts, calyx and corolla inPharbitis nil

The structure and ontogeny of the foliage leaves, bracts, bracteoles, calyx and corolla ofPharbitis nil were investigated, with special reference to the development of the lamina and the procambium. Reproductive organs used are those of a terminal inflorescence and axillary flowers induced by a single 16 hr dark period given to the seedling. The foliage leaf consists of the petiole and the broad lamina. Bracts show various forms and structures, which fluctuate from a lower leafy bract to an upper scaly one in a terminal inflorescence. The sepal is scaly. The corolla is funnel-shaped, and composed of five wedge-shaped petals. In the lamina of the foliage leaf primordium, marginal growth is followed by active growth by the plate meristem, and procambial strands of lateral veins differentiate from the residual meristem. The primordium of the lowest bract of the terminal inflorescence has already been initiated before the dark period, and develops into the bract, the residual meristem disappearing after the treatment. The leafy bract shows marginal growth and growth by the plate meristem similar to that of the foliage leaf, but of short duration. The activity of marginal growth of the scaly bract and the sepal decreases rapidly and procambial strands of lateral veins differentiate acropetally from highly vacuolated cells. The activity of marginal growth of the petal decreases gradually, and derivatives of the marginal meristem divide as a plate meristem. The corolla tube is initiated by co-operation of interprimordial growth and marginal growth of petal primordia.     

马先蒿属花冠无喙类的花器官发生

对花冠无喙类密穗马先蒿(Pedicularis densispica)和大王马先蒿(P．rex)的花器官电镜扫描发现,两种不同花冠型(无齿和具齿)的马先蒿花部器官发生和发育初期十分相似,表现为明显的单轴对称。2个萼片原基首先发生于花顶的近轴侧位,然后沿花顶边缘向远轴端发育形成--马蹄形结构。密穗马先蒿在近轴中部又出现1枚萼片原基,随后马蹄形结构分化出4枚萼片,并与近轴中部的原基愈合后构成5齿萼片;而大王马先蒿的2齿萼片直接由马蹄形结构发育而成。5枚独立的花瓣原基随后发生,但发育相对滞后;除近轴中部位置1枚空缺外,4枚雄蕊原基与花瓣原基位置呈交互发生;2个心皮原基同时在拱形花顶的近轴和远轴端发生,剩余的花顶形成中间的隔膜,并与2个心皮形成中轴胎座。对马先蒿与金鱼草(Antirrhinum majus)和毛地黄(Digitalis purpurea)花器官发生和发育初期的特征进行了比较,讨论了马先蒿属花冠对称性变化的意义。     

Pyrophosphorylases in Solanum tuberosum: I. Changes in ADP-Glucose and UDP-Glucose Pyrophosphorylase Activities Associated with Starch Biosynthesis during Tuberization, Maturation, and Storage of Potatoes

Changes in ADP-glucose and UDP-glucose pyrophosphorylase activities were followed during tuber development of Solanum tuberosum and prolonged storage at 4 and 11 C. Potato tuberization was accompanied by a sharp increase in starch synthesis simultaneous with a marked rise in ADP-glucose pyrophosphorylase activity. When tubers reached an average diameter of 1 centimeter (0.5 gram average tuber weight) and had already established 58% starch on a dry weight basis, ADP-glucose pyrophosphorylase increased 16- to 24-fold over its activity seen in low starch containing stolon tissue. During this same period UDP-glucose pyrophosphorylase increased approximately 2- to 3-fold. Although participation of UDP-glucose in starch formation can not be neglected, it is suggested that the onset of rapid non-photosynthetic potato tuber starch biosynthesis may be closely related to the simultaneous increase in ADP-glucose pyrophosphorylase activity.     

The different large subunit isoforms of Arabidopsis thaliana ADP-glucose pyrophosphorylase confer distinct kinetic and regulatory properties to the heterotetrameric enzyme

ADP-glucose pyrophosphorylase catalyzes the first and limiting step in starch biosynthesis and is allosterically regulated by the levels of 3-phosphoglycerate and phosphate in plants. ADP-glucose pyrophosphorylases from plants are heterotetramers composed of two types of subunits (small and large). In this study, the six Arabidopsis thaliana genes coding for ADP-glucose pyrophosphorylase isoforms (two small and four large subunits) have been cloned and expressed in an Escherichia coli mutant deficient in ADP-glucose pyrophosphorylase activity. The co-expression of the small subunit APS1 with the different Arabidopsis large subunits (APL1, APL2, APL3, and APL4) resulted in heterotetramers with different regulatory and kinetic properties. Heterotetramers composed of APS1 and APL1 showed the highest sensitivity to the allosteric effectors as well as the highest apparent affinity for the substrates (glucose-1-phosphate and ATP), whereas heterotetramers formed by APS1 and APL2 showed the lower response to allosteric effectors and the lower affinity for the substrates. No activity was detected for the second gene coding for a small subunit isoform (APS2) annotated in the Arabidopsis genome. This lack of activity is possibly due to the absence of essential amino acids involved in catalysis and/or in the binding of glucose-1-phosphate and 3-phosphoglycerate. Kinetic and regulatory properties of the different heterotetramers, together with sequence analysis has allowed us to make a distinction between sink and source enzymes, because the combination of different large subunits would provide a high plasticity to ADP-glucose pyrophosphorylase activity and regulation. This is the first experimental data concerning the role that all the ADP-glucose pyrophosphorylase isoforms play in a single plant species. This phenomenon could have an important role in vivo, because different large subunits would confer distinct regulatory properties to ADP-glucose pyrophosphorylase according to the necessities for starch synthesis in a given tissue.     

STRUCTURE AND DEVELOPMENT OF THE PERIANTH IN DOWNINGIA BACIGALUPII

The structure and ontogeny of the calyx and corolla of Downingia bacigalupii Weiler (Campanulaceae; Lobelioideae) were investigated for the purpose of comparing perianth development with previous observations on the floral bract, as well as elucidating the mechanism of development of the zygomorphic, sympetalous corolla. Sepals are uni-traced with a palmate, reticulate venation. They have basal and apical hydathodes, as well as storage tracheids. Sepals show a reduction in size, venation and hydathode number when compared to the bract. The pentamerous, zygomorphic corolla is bilabiate, consisting of a three-lobed adaxial lip and a two-lobed abaxial lip connected by a short tubular region. The constituent petal lobes are also uni-traced and have a reticulate venation, resembling that of the sepal and bract, but lack storage tracheids and hydathodes. Sepals arise in an adaxial to abaxial succession and are initiated in the outer corpus layer of the floral apex. Expansion of the floral apex follows and is accompanied by the establishment of a second tunica layer. Sepals undergo apical, marginal, and intercalary growth accompanied by acropetal differentiation of procambium. The petals arise simultaneously and are initiated in the second tunica layer and the outer corpus cells. After initiation, the petals exhibit a period of apical and marginal growth followed by intercalary growth. Apical growth in petals is less protracted than in sepals, but plate meristem activity is more extensive. The free petal lobes become temporarily fused by an interlocking of marginal epidermal layers, but they separate at anthesis. Zonal growth beneath the originally free lobes forms the tube and lip regions of the sympetalous corolla. Zygomorphy is evident from the time of initiation of petals and is accentuated by later differential growth. Comparative observations of corolla ontogeny in autogamous species of Doumingia indicate that the reduced corollas in these taxa are derived by a simple process of neoteny.     

Theligonum cynocrambe: Developmental morphology of a peculiar rubiaceous herb

The annual Mediterranean herbTheligonum cynocrambe shows a peculiar combination of morphological characters, e.g., switch from decussate to spiral phyllotaxis with 90–100° divergence, combined with a change from interpetiolar to lateral stipules, anemophily, lack of calyx, flowers often dimerous to trimerous, corolla fused in both male and female flowers, male flowers extra-axillary, with 2–19 stamens per flower, female flowers axillary, with inferior uniovulate ovary, basilateral style and perianth, nut-like fruits with elaiosome. In male flowers the androecium emerges as an (uneven) elliptical rim with a central depression. This common girdling primordium is divided up into several stamen primordia. In male flowers with low stamen number the stamen primordia may occupy the corners alternating with the corolla lobes. There are no epipetalous androecial primordia that secondarily divide into stamens. Male flowers occasionally show a hemispherical base that may be interpreted as remnant of the inferior ovary. In female flowers a ring primordium grows into a tube on which the petal lobes arise. The perianth and style become displaced adaxially by uneven growth of the inferior ovary. The ovary is basically bilocular. The lower region of the ovary is provided with a septum that is overtopped and hidden by the single curved ovule.Theligonum is referred to theRubiaceae-Rubioideae, with theAnthospermeae andPaederieae as most closely related tribes.     

红肉脐橙前期花芽形态分化研究

为了摸清红肉脐橙的花芽分化时期,以便在适宜的时期采取措施调控花芽分化,确保每年都有适宜的花量,为高产稳产奠定基础。2006～2007年,笔者采用石蜡切片法观察了红肉脐橙花芽形态分化过程。结果表明,红肉脐橙花芽分化从11月上旬开始,11月下旬开始萼片分化,翌年1月中旬进入花瓣分化期,2月上旬雄蕊、雌蕊分化开始,每个时期都历时较长。其过程可分为生理分化期、花芽分化期、花萼分化期、花瓣分化期、雄蕊分化期、雌蕊分化期和子房形成期7个时期。红肉脐橙花量大,其花芽分化过程比较缓慢,分化期也较分散,分化时间长,每个时期都有重叠交叉现象。     

Translocation of 14C-sucrose in Relation to Changes in Carbohydrate Content in Rose Corollas Cut at Different Stages of Development

The dry matter and carbohydrate contents of intact growing ‘Sonia’rose corollas were measured from an immature bud to full expansionof the petals. Reducing sugars and starch, but not sucrose,accumulated throughout most of the corolla development. Thesefindings were compared with the carbohydrate changes in thecorollas of flowers cut at different stages and allowed to agewith their stems either in water or in a sucrose-containingsolution. For a few days after cutting the carbohydrate metabolismof the cut flower roughly paralleled that of the intact floweruntil starch hydrolysed to maintain the soluble carbohydratepool. Feeding with the sucrose solution maintained the solublecarbohydrate levels and retarded the hydrolysis of starch. The cut flowers were fed with ¹⁴C-sucrose and the labelled metabolitesin the leaves and flowers were analysed. Active incorporationof ¹⁴C into ethanol-soluble carbohydrates, starch and ethanol-insolublematerial was found indicating that an active anabolic phaseprecedes the catabolic phase during the senescence of the cutflower. The findings are discussed in relation to the source-sinkhypothesis of flower development, with regard to the senescenceand growth of the corollas of cut and intact flowers respectively.