Floral rewards and pollination in Cytiseae (Fabaceae)

 Cytiseae have been reported to be mostly nectar-lacking, yet some taxa secrete nectar from extrastaminal nectaries. We studied the pollination biology of four shrubby species of Cytiseae (Cytisophyllum sessilifolium (L.) Lang, Spartium junceum L., Genista radiata (L.) Scop., Genista cilentina Valsecchi) which differ for ecology, distribution and population size. All species resulted obliged xenogamous, insect visits being necessary for successful pollination. Bumblebees and solitary bees are the principal pollinators, but also many beetles, some hover-flies, and few bugs visit flowers. Pollinator specificity is low, and this may be the reason of the scarce seed set compared to the number of ovules. Pollen is the main reward, but traces of glucose were detected in all species, at the base of vexillum or on the reproductive column. Nectar production is irregular in time, and apparently unpredictable. We suppose that nectar may play a role in attracting pollinators determining their right position for a successful pollination. Received August 12, 2002; accepted November 25, 2002 Published online: June 2, 2003     

Floral organogenesis of Chloranthus sessilifolius, with special emphasis on the morphological nature of the androecium of Chloranthus (Chloranthaceae)

 Floral organogenesis of Chloranthus sessilifolius K. F. Wu is described. The inflorescence primordium is dome-like in the beginning and then elongates, and bract primordia initiate almost decussately. Each floral primordium, arising from the axil of a bract, soon becomes a scale-like structure, with three primordia of androecial lobes originating from its abaxial part, and the gynoecial primordium in adaxial position. As the androecial lobes become more distinct, four thecae are already in differentiation, and the gynoecial primordium appears as a shallow disc. The androecial lobes do not extend their length until the thecae approach maturity and the stigma is differentiated. The androecial lobes are united at all the stages of development, and the entire androecium falls off as a unit at the end of anthesis. Based on these results, combined with published evidence from neobotany, palaeobotany and phylogenetic studies, the morphological nature of the androecium of Chloranthus is further discussed. Our studies support the viewpoint that the androecial structure of Chloranthus may have arisen by splitting of a single stamen with 2 marginal thecae. Received May 2, 2001 Accepted December 18, 2001     

Nectar biodiversity: a short review

 Nectaries differ in many aspects but a common feature is some kind of advantage for the plant conferred by foraging of consumers which may defend the plant from predators in the case of extrafloral nectaries, or be agents of pollination in the case of floral nectaries. This minireview is concerned mainly with floral nectaries and examines the following characteristics: position in flower; nectary structure; origin of carbohydrates, aminoacids and proteins; manner of exposure of nectar; site of nectar presentation; volume and production of nectar in time; sexual expression of flower and nectary morphology; nectar composition and floral sexual expression; variability of nectar composition; fate of nectar; energy cost of nectar production. The species of certain large families, such as Brassicaceae, Lamiaceae and Asteraceae, resemble each other in nectary organisation; other families, such as Cucurbitaceae and Ranunculaceae, have various types of organisation. A scheme is presented to illustrate factors influencing nectary and nectar biodiversity. Received July 23, 2002; accepted September 18, 2002 Published online: June 2, 2003     

Floral nectaries and nectar production in brown mustard (Brassica juncea) and white mustard (Sinapis alba) (Brassicaceae)

 The dynamics and abundance of nectar secretion as well as sugar productivity were studied in flowers of brown mustard (Brassica juncea) cv. Małopolska and white mustard (Sinapis alba) cv. Borowska. Moreover, floral nectaries were examined under LM and SEM. In both cultivars lateral and median pairs of nectaries secreted nectar. However, differences were found in morphology and activity of these pairs. The lateral nectaries produced more nectar than the median ones. Nectar secretion started at loose bud and peaked during anther dehiscence. Average amount of nectarsecreted by 100 flowers of cv. Małopolska and cv. Borowska were 119.9 mg and 134 mg. Mean concentration of nectar was 26.7% and 23.4%, respectively. One hundred flowers of cv. Małopolska and cv. Borowska secreted 28.4 mg and 24.9 mg of sugars in nectar. Estimated sugar productivity per 1 ha of crop was 65.5 kg and 71.2 kg, respectively. Received August 28, 2002; accepted November 2, 2002 Published online: June 2, 2003     

Floral morphology and reproductive success in the orchid Epipactis helleborine: regional and local across-habitat variation

 The terrestrial orchid Epipactis helleborine is a morphologically variable species with a wide distribution in Europe. It is pollinated by social wasps, and most populations show the morphological characteristics of outcrossing species. However, local predominantly selfing subspecies and varieties have been documented from drier habitats. To document geographic variation in floral morphology, ability to produce seeds through autogamy, and reproductive success in E. helleborine, we sampled 13 populations from three geographic regions along a latitudinal gradient of c. 1000 km from northern to southern Sweden. In the southernmost region, populations in dry and mesic habitats were compared. Supplemental hand-pollination was conducted to determine whether among-population variation in fruit set could be explained by differences in the natural level of pollination, and whether any relationship between floral morphology and fruit production could be explained by interactions with pollinators. Bagging experiments showed no evidence of autogamy in any of the study populations. Number of flowers, pollinia removal and fruit set varied significantly among populations but did not differ among regions. Pollinia removal was positively correlated with population size and both pollinia removal and fruit set were lower in dry than in mesic habitats. At the level of the individual plant, the number of pollinia removed increased more rapidly with flower number than did number of fruits produced. The hand-pollination experiment indicated that the positive relationship between number of flowers and fruit production was due to a higher degree of pollen limitation in plants with few flowers than in plants with many flowers. The experiment also showed that variation in the level of pollen limitation could only partly explain variation in fruit set among populations. Received November 6, 2001; accepted April 27, 2002 Published online: December 3, 2002     

Effect of rootstocks on floral nectar composition in apple cultivars

 Insect attraction of flowers in apple cultivars is an essential question from the viewpoint of fruit yield, since apple flowers are autosterile and entomogamous. Therefore, it is important to study the two most significant factors of primary attraction, pollen and nectar production. The composition of nectar in apple cultivars on different rootstocks was investigated, putting an emphasis on sugar components. It was stated that the composition of nectar and the ratio of mono- and disaccharides was almost identical and constant in trees on rootstock M.4, whereas on other rootstocks the effects of the season were more pronounced. In triploid cultivars on rootstocks M.9 and M.26 nectar can be characterised by the dominance of sucrose. Relationship was found between free fertilisation rate of cultivars and sugar amount of nectar. In triploid cultivars nectar sugar content was usually lower than in diploid ones, and reached the bee visitation threshold only in some years, influenced also by the effect of rootstocks. Received August 27, 2002; accepted November 16, 2002 Published online: June 2, 2003     

Female flowers and inflorescences of Didymelaceae

 In molecular analyses Didymelaceae together with Buxaceae form a fairly well-supported clade among families near the base of eudicots. Only little is known, however, about the flowers and inflorescences of Didymelaceae. In this study, the structure of the female flowers and inflorescences of Didymeles integrifolia was studied. Flowers are unicarpellate and orientation of the carpel is slightly deflected abaxially as in Proteaceae. Otherwise, Didymelaceae share many features of the gynoecium with Buxaceae and some other basal eudicots: the carpels are ascidiate in the lower half; anthetic carpels are completely closed by postgenital fusion; stigma is double-crested and widely decurrent; stigmatic papillae are unicellular and pear-shaped; the pollen tube transmitting tract is extensive and prominently differentiated; fruits are fleshy drupes with persistent stigma and style. However, the exceedingly elongate base of the integuments of Didymelaceae is an unusual feature among basal eudicots and even angiosperms. Received October 31, 2002; accepted December 17, 2002 Published online: March 31, 2003     

The correlation between development of atypical bisexual flowers and phylogeny in the Aroideae (Araceae)

 In the intermediate zone of the inflorescence of genera of Aroideae one can find flowers with male and female characteristics. Until now, two types of developmental sequences of atypical bisexual flowers (ABFs) have been recognized: the Philodendron type and the Cercestis type. In the Philodendron type, bisexual flowers generally consist of functional carpels and staminodes inserted on the same whorl. In the Cercestis type, the gynoecium and stamens are inserted on two different whorls. These different ontogenetic patterns represent two different pathways in the evolution of unisexual flowers in this subfamily. A molecular phylogenetic analysis of 33 genera of Araceae, based on the chloroplast trnL intron and trnL–F intergenic spacer sequences was carried out. We use this phylogenetic analysis and those published by French et al. (1995) and Mayo et al. (1997) to examine the distribution of the two types of ABFs in selected genera. Our results suggest that the two developmental patterns of ABFs in Aroideae sensu Mayo et al. (1997) do not correspond to two separate evolutionary lineages but rather are more or less consistent within clades. Although this new molecular phylogeny does not include all aroid genera, it corroborates in general, at the subfamily level, the molecular analysis of French et al. (1995) based on chloroplast DNA restriction site data and the analysis of Mayo et al. (1997) based on morphological and anatomical data. Received March 15, 2001 Accepted October 11, 2001     

Inflorescence development of Whytockia (Epithemateae, Gesneriaceae) and phylogenetic implications within Gesneriaceae

 The inflorescence development in Whytockia has been studied in order to explore the developmental basis for inflorescence architecture. The developmental pattern of the pair-flowered cyme in Whytockia basically conforms to that of most members in Gesneriaceae. However, the additional flower beside the terminal one in Whytockia is not equivalent to the frontal flower as in other Gesneriaceae because the former is located in the front-lateral position while the latter is in the front-median position. Also, the zigzag monochasial branching system in Whytockia represents the consecutive front-lateral branches rather than the lateral branches as in other Gesneriaceae. The inflorescence in Whytockia is flowering in a basipetal sequence, and its seemingly acropetal flowering sequence is due to the vigorous development of the consecutive front-lateral branches. In addition, the inflorescence of Whytockia does not represent the basic unit of the inflorescence in Epithemateae, and it is derived as compared to that of Rhynchoglossum. The development relationships of the inflorescence between Whytockia and its allies in Epithemateae are discussed on the basis of developmental and comparative evidence. Received February 15, 2002; accepted September 17, 2002 Published online: December 11, 2002     

Floral structure and development in the monoecious palm Gaussia attenuata (Arecaceae; Arecoideae)

Background and Aims

Sexual dimorphism, at both the flower and plant level, is widespread in the palm family (Arecaceae), in contrast to the situation in angiosperms as a whole. The tribe Chamaedoreeae is of special interest for studies of the evolution of sexual expression since dioecy appears to have evolved independently twice in this group from a monoecious ancestor. In order to understand the underlying evolutionary pathways, it is important to obtain detailed information on flower structure and development in each of the main clades.

Methods

Dissection and light and scanning electron microscopy were performed on developing flowers of Gaussia attenuata, a neotropical species belonging to one of the three monoecious genera of the tribe.

Key Results

Like species of the other monoecious genera of the Chamaedoreeae (namely Hyophorbe and Synechanthus), G. attenuata produces a bisexual flower cluster known as an acervulus, consisting of a row of male flowers with a basal female flower. Whereas the sterile androecium of female flowers terminated its development at an early stage of floral ontogeny, the pistillode of male flowers was large in size but with no recognizable ovule, developing for a longer period of time. Conspicuous nectary differentiation in the pistillode suggested a possible role in pollinator attraction.

Conclusions

Gaussia attenuata displays a number of floral characters that are likely to be ancestral to the tribe, notably the acervulus flower cluster, which is conserved in the other monoecious genera and also (albeit in a unisexual male form) in the dioecious genera (Wendlandiella and a few species of Chamaedorea). Comparison with earlier data from other genera suggests that large nectariferous pistillodes and early arrest in staminode development might also be regarded as ancestral characters in this tribe.     

Floral nectary ultrastructure of Prunus persica (L.) Batch cv. Forastero (Newcomer), an Argentine peach

 Flowers of Prunus persica (L.) Batch. cv. Forastero have an orange toral nectary. The nectariferous tissue was formed by densely packed parenchyma cells (secretory cells) and an epidermis with hairs and modified stomata. The epidermal cells were highly vacuolated with a striated cuticle. The ultrastructure of these cells contained a cytoplasm with endoplasmic reticulum, plastids, mitochondria and dictyosomes. Sub-epidermal cells were barely vacuolated and their ultrastructure was similar to that of the epidermal cells. Differences were observed only in the endoplasmic reticulum, which is organized in a parallel configuration. Plasmodesmata were found between adjacent secretory cells and between secretory and epidermal cells. An electron dense secretion occurred in the intercellular spaces and between the external tangential wall and the cuticle of the epidermal cells. According to the ultrastructural observations, the sugar solution could be passed through the symplast or the apoplast. The nectar could be exuded from the stomata and the micro-channels of the cuticle covering the epidermal cells. Received July 7, 2002; accepted September 24, 2002 Published online: June 2, 2003     

Approach to the early sporangial development in angiosperms considering meiosis control and cellular differentiation

 Applying current data on cell differentia- tion and meiosis control to the early sporangial development in angiosperms, a strict relationship between cell lineage and its differentiation fate is rejected. An evaluation of cytological features indicative of a meiotic (sporogenous) fate discards the sterilization phenomenon and introduces the premeiotic cellular differentiation (PCD) concept. The early sporangial development comprises 5 basic steps and 4 cellular stages, where PCD and meiosis extension and gradient are related to mechanisms of spore mother cell selection. Concepts here discussed explain the exceptions to the normal early sporangial development and allow a precise definition of archesporium and archespore. PCD and meiotic extension and gradient recover more information of the early sporangial development, distinguishing different developmental patterns leading to the same final result and retaining slight developmental differences. However, there are no early developmental characteristics distinctive of andro- or gynosporangia. Therefore, the heterosporic condition is not related to early developmental changes. Received January 5, 2001 Accepted August 29, 2001     

Floral anatomy of Bromeliaceae, with particular reference to the evolution of epigyny and septal nectaries in commelinid monocots

Floral anatomy is described in ten genera of Bromeliaceae, including three members of subfamily Bromelioideae, three Tillandsioideae, and four genera of the polyphyletic subfamily Pitcairnioideae (including Brocchinia, the putatively basal genus of Bromeliaceae). Bromeliaceae are probably unique in the order Poales in possessing septal nectaries and epigynous or semi-epigynous flowers. Evidence presented here from floral ontogeny, vasculature, and the relative positions of nectary and ovules indicates that there could have been one or more reversals to apparent hypogyny in Bromeliaceae, although this hypothesis requires a better-resolved phylogeny. Such evolutionary reversals probably evolved in response to specialist pollinators, and in conjunction with other aspects of floral morphology of Bromeliaceae, such as the petal appendages of some species. The ovary is initiated in an inferior position even in semi-epigynous or hypogynous species. The ovary of all so-called hypogynous Bromeliaceae is actually semi-inferior, because the septal nectary is infralocular; in these species the nectaries have a labyrinthine surface and many vascular bundles. Brocchinia differs from most other fully epigynous species in that each carpel is secretory at the apex and reproductive, rather than secretory, at the base.     

Oxypetalum banksii subsp. banksii: a taxon of Asclepiadaceae with an extragynoecial compitum

 The floral morphology of seven Oxypetalum species and, in particular, the spatial relationship between the five stigmatic chambers and two separate ovaries of their flowers with respect to transmission of the pollen tube are studied. In all species, except O. banksii subsp. banksii, floral morphology is similar to that in other Asclepiadeae, and the flowers pollinated with one pollinium develop only one follicle, which means compitum absence. In O. banksii subsp. banksii flowers, the secretory interstaminal tissue lines the inner walls of the stigmatic chambers as in the other species studied, but it also reaches the upper part of the inner surface of the filament tube, where it surrounds the styles, an unprecedented feature for Asclepiadaceae. This tissue secretes nectar and mucilage; the latter acts as transmitting medium for the growth of pollen tubes from pollinia inserted and hydrated in stigmatic chambers (“hyperstigmas”). Mucilage also functions as an extragynoecial compitum: in flowers pollinated with one pollinium both carpels develop into a follicle. Received August 28, 2001; accepted April 9, 2002 Published online: October 14, 2002 Addresses of the authors: Milene Faria Vieira (e-mail: mfvieira@mail.ufv.br), Departamento de Biologia Vegetal, Universidade Federal de Vi?osa, 36571-000, Vi?osa, Minas Gerais, Brazil. George John Shepherd, Departamento de Botanica, Instituto de Biologia, Universidade Estadual de Campinas, C.P. 6109, 13083-970, Campinas, S?o Paulo, Brazil.     

Iridoid glucosides of the genus Veronica s.l. and their systematic significance

 The mode of reproduction, pollen production, chromosome numbers, genetic variation (RAPD, allozymes) and overall similarity were studied in 6 species of Hieracium sect. Alpina in the Tatry Mts. (the Western Carpathians, Slovakia). All species were confirmed to be agamospermous and, except of H. krivanense and H. slovacum, lacking pollen grains. For the first time, a chromosome number is reported for H. krivanense (2n=4x=36). Considerable genetic variation was revealed in H. alpinum and a correlation between geographic and genetic distances was found in this species. Between-population variation in RAPD and allozyme phenotypes was found in H. pinetophilum and H. crassipedipilum. In all other species, allozyme and RAPD variation was low or absent. With few exceptions, the species differ in their allozyme as well as RAPD patterns. The relatedness of one population of endemic H. slovacum and H. halleri was confirmed. It is shown, that Carpathian species of the H. fritzei group are derived from at least two ancestors. Received July 3, 2000; accepted June 24, 2002 Published online: November 20, 2002 Addresses of the authors: H. Štorchová, (e-mail: storchova@ueb.cas.cz) Institute of Experimental Botany, Academy of Sciences of the Czech Republic, Rozvojová 135, CZ-165 00 Praha 6, Czech Republic. I. V. Bartish, J. Chrtek Jr., J. Kirschner, M. Tetera, J. Štěpánek, Institute of Botany, Academy of Sciences of the Czech Republic, CZ-252 43 Pruhonice, Czech Republic.     

Tests of the accuracy of cladograms in Gilia (Polemoniaceae) and some other angiosperm genera

 This paper deals with the use of cladistic methods and cladograms in phylogeny reconstruction in plant groups containing numerous taxa. How accurate are the cladograms as to details? Accuracy tests at the level of details require an independently known phylogeny, which excludes most plant groups, but such tests can be carried out in domesticated and experimental plant groups which have documented pedigrees. Four such tests are known and are presented here: a new case in Gilia and three previously published cases in Avena, Hordeum, and Helianthus. The four cases include domesticated and experimental plants, use of morphological and molecular evidence, and presence of dichotomous as well as reticulate phylogenies. The cladograms of the four plant groups all differ in significant details from the known pedigrees. These results are discussed in relation to problems of interpretation of cladograms. Received March 21, 2000 Accepted August 16, 2001     

Influence of elevated CO2 and ultraviolet-B radiation levels on floral nectar production: a nectary-morphological perspective

 Investigations of the effects of two global events – elevated CO₂ levels and enhanced ultraviolet-B (UV-B) radiation – on floral nectar production are reviewed from twelve dicotyledonous families. Furthermore, to allow comparisons between nectary morphology and nectar production in treated plants of these fifteen species, new data on floral nectary structure are provided for Malcolmia maritima (L.) R. Br. (Brassicaceae) and Scabiosa columbaria L. (Dipsacaceae). All but the last taxon possessed mesenchymatic floral nectaries with surface stomata. Few clear relationships existed between nectary morphology and various physiological responses to CO₂ or UV-B enrichment, indicating that species responded notwithstanding nectary structure itself. Overall, nectar-solute concentration was least affected by elevated CO₂ or UV-B radiation; consequently, changes in nectar volume were responsible for differences in nectar-sugar production per flower. Three species of Fabaceae experienced no change in floral nectar production upon exposure to elevated CO₂. To date, no study of enhanced UV-B radiation reported a consistent reduction in floral nectar production; three species of Brassicaceae responded differently, but various levels of ozone depletion were simulated. Experimentation with more taxa – including those possessing nectary types such as septal (gynopleural) nectaries (e.g. many monocotyledons) or aggregations of glandular trichomes – and expanding such physiological studies to species possessing extrafloral nectaries, are recommended. Received August 8, 2002; accepted November 23, 2002 Published online: June 2, 2003     

Floral ontogeny and systematic position of the Didiereaceae

Floral ontogenetical data from all four genera of the Didiereaceae (s.str.) are presented for the first time. All Didiereaceae s.str. are dioecious, having unisexual flowers with organ rudiments of the opposite sex. Two median bracts followed by a tetramerous perianth (two alternating dimerous ``whorls'), a slightly complex androecium with 6–12 stamens in a single row (on a common ring primordium), four of which mostly alternating with the perianth members, and one basal ovule connecting three free septa at their very base are flower characters in Didiereaceae, supporting phylogenetic analyses based on nucleotide sequence data. Closest relatives are the (formerly) portulacaceous genera Portulacaria (5 stamens alternating with the perianth), Ceraria (5 stamens alternating with the perianth), and Calyptrotheca (many stamens), all with pentamerous perianths, from which the tetramerous perianth in Didiereaceae can be derived. Applequist and Wallace (2003) included these three genera in an expanded family Didiereaceae (with three subfamilies).     

Floral ontogeny and anatomy inKoelreuteria with special emphasis on monosymmetry and septal cavities

The floral ontogeny and anatomy ofKoelreuteria paniculata have been investigated to understand the developmental basis for the occurring monosymmetry and the origin of the septal cavities. Petals arise sequentially and one petal is missing between sepals 3 and 5, or rarely between sepals 2 and 5. The eight stamens arise sequentially before petal initiation is completed. The last formed petal and one stamen arise on a common primordium. Two stamen positions are empty (opposite the petal between the sepals 2 and 5, and the petal between sepal 1 and 3); consequently two antesepalous stamens have become displaced. The derivation of octandry from a diplostemonous ancestry, and reduction of the petal are discussed. The triangular gynoecium has a strong impact in obliquely reorganizing the symmetry of the flower, loss of organs, and shifts of stamens. The so-called septal slits occurring within the style are a deepreaching non-nectariferous extension of the stigma. Alternating locular furrows are present which could play a role as pollen transmitting tissue and in the loculicid dehiscence of the capsule.     

Floral development of an asexual and female-like mutant carrying two deletions in gynoecium-suppressing and stamen-promoting functional regions on the Y chromosome of the dioecious plant Silene latifolia

Sexual dimorphism is controlled by genes on the Y chromosome in the dioecious plant Silene latifolia. K034 is the first mutant with female flowers and asexual flowers in one individual. Its stamens are suppressed completely, and its gynoecium exhibits two suppression patterns. One gynoecium resembles a thin rod, as in wild-type males (asexual flower); the other is imperfectly suppressed, having 1-3 carpels (female-like flower). The ratio of these patterns was 9 : 1. To exclude the possibility of chimerism in K034, we crossed a female-like flower of K034 with a wild-type male. Progeny obtained from this crossing had asexual and female-like flowers in one individual. This two-flower-type phenotype was inherited without separating. To examine the identity of flower organs in K034, we analyzed the development of asexual and female-like flowers using scanning electron microscopy and in situ hybridization with SLM1 and SLM2 (orthologs of AGAMOUS and PISTILLATA, respectively) as probes. Mitotic spreads of root tip chromosomes from hairy root cultures showed that K034 had 25 chromosomes. Fluorescent in situ hybridization analysis, using a subtelomeric repetitive sequence (KpnI subfamily) as a probe, indicated that K034 possessed two X chromosomes and one Y chromosome (Y(d)), of which Y(d) had been rearranged to lose the pseudoautosomal region (PAR). PCR analysis using Y-specific sequence-tagged site (STS) markers clarified that Y(d) of K034 had two other deletions in gynoecium-suppressing and stamen-promoting regions. It is reasonable to suggest that these sex chromosomal abnormalities resulted in two abnormal sexual phenotypes: the asexual and imperfect female (female-like) flowers in K034.