Floral biology of Nymphaea mexicana zucc. (Nymphaeaceae)

The floral biology of Nymphaea mexicana Zuccarini has been investigated. Anthesis is diurnal with flowers opening and closing for two consecutive days. Flowers on the first day of anthesis are functionally female with the stigmatic papillae secreting a small quantity of fluid. Total dissolved solids of the stigmatic fluid range from 3 to 4%; glucose and fructose are in equal concentrations and a large number of free amino acids are present. Flowers are primarily visited by Dialictus bees, flies and beetles. These insects typically land on the wet stigmatic surface where the secretion loosens pollen from the insects' body. On the second day of anthesis, the anthers dehisce and the stigmatic fluid disappears. This latter phenomenon is associated with a loss of stigmatic receptivity. Since protogyny is complete, flowers are obligatorily entomophilous. After anthesis, the flower closes and submerges. The perianth and androecium abscise and decompose as the fruit matures. The fruit yields large trichomic seeds. The phylogenetic position of N. mexicana within the genus is discussed.     

A note on the pollination of Nymphaea amazonum Mart. & Zucc. (Nymphaeaceae)

Nymphaea amazonum Mart. & Zucc. a nocturnally flowering species, is visited by the scarab beetleCyclocephala mollis Endrödi in the Pantanal region of Mato Grosso, Brazil. Beetles were found in all flowers examined and some remain in the flowers during the day. The beetles are dusted with pollen in second-day flowers and carry it to first-day flowers which have receptive stigmas.     

Reproductive biology of Nymphaea capensis Thunb. var. zanxibariensis (Casp.) Verdc. (Nymphaeaceae)

Anthesis in Nymphaea capensis var. zanzibariensis is diurnal with flowers opening and closing for three consecutive days. On the first day of anthesis, the stigmatic papillae secrete fluid and the outermost anthers are dehiscent. On the second day of anthesis the stamens form a cone above the dry stigmatic cup. The middle stamens open and turn outward. On the third day of flowering, all the stamens open and the dry stigmatic cup is exposed. The flowers are homogamous and not protogynous as the other Nymphaea. The gynoecium of the self-compatible N. capensis var. zanzibariensis , is characterized by a wet papillate stigma, a short hollow style, and secretory cells on the ventral surface of the ovary. The pollen is released on the receptive stigma. Following initial growth in intercellular spaces in the transmitting tract of the stigma, pollen tubes travel through the stylar canal and into the ovary.     

Germination requirements and seed banks of some nymphaeid macrophytes: Nymphaea alba L., Nuphar lutea (L.) Sm. and Nymphoides peltata (Gmel.) O. Kuntze

SUMMARY.

• 1 Germination experiments demonstrated that the innate dormancy of the seeds of Nymphaea alba L., Nuphar lutea (L.) Sm. and Nymphoides peltata (Gmel.) O. Kuntze could be overcome by a cold treatment. Light stimulated the germination of the three species. Hypoxic conditions stimulated the germination of Nymphaea alba and Nuphar lutea seeds but the seeds of Nymphoides peltata did not germinate under these conditions.
• 2 Experimental seed banks of Nymphaea alba, Nuphar lutea and Nymphoides peltata were laid out in three water bodies, varying in pH and alkalinity. Germination patterns indicated that Nymphaea alba and Nuphar lutea produce transient seed banks, but that Nymphoides peliata produces a persistent seed bank. Sampling of natural seed banks and subsequent germination tests were in concordance with the results of the seed bank experiment.
• 3 The experimental above-ground seed banks of Nymphaea alba, Nuphar lutea and Nymphoides peltata showed similar germination patterns in the three selected water bodies, despite the differences in pH and alkalinity between them. However, the distribution of Nymphoides peliata is restricted to well-buffered waters, so that its absence from soft and acid water bodies must be due to post-germination mechanisms and/or processes.
• 4 In aquatic systems where Nymphoides peltata co-exists with the other nymphaeid species studied, it is largely restricted to a bell between the helophytes and the vegetation at deeper sites. The deeper sites were dominated by Nuphar lutea and Nymphaea alba. Germination requirements and seedling emergence from buried seeds of Nymphaea alba, Nuphar lutea and Nymphoides peliata play an important role in the establishment of the zonation pattern of these nymphaeid macrophytes.

     

Developmental morphology of branching flowers in Nymphaea prolifera

Nymphaea and Nuphar (Nymphaeaceae) share an extra-axillary mode of floral inception in the shoot apical meristem (SAM). Some leaf sites along the ontogenetic spiral are occupied by floral primordia lacking a subtending bract. This pattern of flower initiation in leaf sites is repeated inside branching flowers of Nymphaea prolifera (Central and South America). Instead of fertile flowers this species usually produces sterile tuberiferous flowers that act as vegetative propagules. N. prolifera changes the meristem identity from reproductive to vegetative or vice versa repeatedly. Each branching flower first produces some perianth-like leaves, then it switches back to the vegetative meristem identity of the SAM with the formation of foliage leaves and another set of branching flowers. This process is repeated up to three times giving rise to more than 100 vegetative propagules. The developmental morphology of the branching flowers of N. prolifera is described using both microtome sections and scanning electron microscopy.     

Nymphaea tetragona and Nymphaea leibergii (Nymphaeaceae): two species of diminutive water-lilies in North America

Although all past floras of northern North America have recognized only one species of diminutive water-lily, actually two distinct species occur in the region. The circumboreal Nymphaea tetragona Georgi is largely confined to the northwestern part of the continent, extending as far east as Manitoba and occurring in the contiguous United States only in northwest Washington. A second species, N. leibergii Morong, is restricted to, but widely distributed in, northern North America, though it is absent from the extreme northwest. The two species are distinguishable from each other by several floral and foliar characters and the two species form a section, Nymphaea subg. Nymphaea sects. Chamaenymphaea, stat: nov. This is distinguished from the two other sections of subg. Nymphaea in a key, and Nymphaea subg. Nymphaea sect. Xanthantha, stat. nov., is proposed. Sect. Chamaenymphaea is described for North America, synonymy and nomenclature for both species are discussed, and a key including N. odorata Aiton is presented. A lectotype is provided for N. leibergii and the typification of N. tetragona is examined.     

Pollen tube growth and the pollen‐tube pathway of Nymphaea odorata (Nymphaeaceae)

An important aspect of the evolution of carpel closure, or angiospermy, is the relationship between pollen tube growth patterns and internalization of the pollen‐tube pathway. True carpel closure, involving postgenital fusion of inner carpel margins, is inferred to have arisen once within the ancient order Nymphaeales, in the common ancestor of Nymphaeaceae. We studied pollen tube development, from pollination to fertilization, in a natural population of Nymphaea odorata, using hand pollinations and timed flower collections. Pollen germinates in stigmatic secretions within 15 min and pollen tubes enter subdermal transmitting tissue within an hour, following wide intercellular spaces towards the zone of postgenital fusion. At the zone of fusion they turn downwards to grow in narrow spaces between interlocked cells and then wander freely to ovules within ovarian secretions. The pollen‐tube pathway is 2–6 mm long and upper ovules are first penetrated 2.5 h after pollination. Pollen tubes grow at rates of approximately 1 mm/h whether in stigmatic fluid, transmitting tissues or ovarian secretions. Pollen‐tube pathways are structurally diverse across Nymphaeales, yet their pollen tubes have similar morphologies and rapid growth rates. This pattern suggests pollen tube growth innovations preceded and were essential for the evolution of complete carpel closure. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162 , 581–593.     

Isolation and antimicrobial activities of fungi derived from Nymphaea lotus and Nymphaea stellata

This study aimed to isolate fungi from leaves, petals, stalks and stamens of Nymphaea lotus and Nymphaea stellata and explore their bioactive potential with respect to antimicrobial activity and their cytotoxicity on human cell lines. The totals of 210 fungal isolates were obtained from 640 segments. The highest isolation rate was found in petal segments (0.44), while stamens yielded the lowest isolation rate (0.13). One hundred and ninety–five fungal extracts were evaluated for their efficiency against ten pathogenic microorganisms by colorimetric broth microdilution tests. Cell hexane extract from Eupenicillium levitum FNL036 (FNL036CH) gave the strongest antifungal activity against Cryptococcus neoformans and Talaromyces marneffei with minimum inhibitory concentration of 0.5?μg/mL and affected cell morphology of the test microorganisms. Moreover, this extract was non–cytotoxic to human embryonic kidney 293 (293T) and human keratinocytes (HaCAT) cell lines with 50% inhibitory concentration (IC₅₀) values above 100?μg/mL. This finding confirmed that fungi derived from these two Nymphaea species were a novel source of bioactive metabolites.     

Development and Structure of the Root Cortex in Caltha palustris L. and Nymphaea odorata Ait.

Structural features of the mature root cortex and its apoplasticpermeability to dyes have been determined for two dicotyledonouswetland plants of differing habitats: Nymphaea odorata, growingrooted in water and mud, and Caltha palustris, growing in temporalwetlands among cattails. In mature roots, movement of the apoplasticdyes, berberine and safranin, into the roots was blocked atthe hypodermis, indicating the presence of an exodermis. A hypodermiswith an exodermis, i.e. Casparian bands in the outermost uniseriatelayer plus suberin lamellae, is present in both species. InN. odorata, hypodermal walls are further modified with cellulosicsecondary walls. Roots of N. odorata and C. palustris have anendodermis with Casparian bands only. A honeycomb aerenchymais produced by differential expansion in N. odorata and includesastrosclereids and diaphragms, while roots of C. palustris haveno aerenchyma, but some irregular lacunae are found in old roots.These aspects of cortex structure are related to an open meristemorganization, with unusual patterns of cell divisions in certainground meristem cells (called semi-regular hexagon cells) ofN. odorata. The correlation between aerenchyma pattern and hypodermalstructure appears to be related to habitat differences.Copyright2000 Annals of Botany Company Caltha palustris, Nymphaea odorata, root development, cortex, endodermis, aerenchyma, exodermis, hypodermis, permeability, wetland plants     

Sexual differentiation in Asparagus officinalis L.

Summary Sexual dimorphism in the dioecious plant Asparagus officinalis L. was examined by two-dimensional (2-D) electrophoresis of both total proteins and newly synthesized proteins from cladophylls (leaves), whole mature flowers and homologous sex organs (i.e. true female ovaries and small sterile ovaries from male flowers). Polypeptides isolated from cladophylls of male and female plants were practically indistinguishable; the flowers, however, showed a distinct set of specific proteins, some of which differed between the two sexes. While the total protein profiles of isolated ovaries from male and female plants were very similar, the patterns were strikingly different after the tissues were pulsed with ³⁵S-methionine: mature male ovaries showed a number of newly synthesized proteins, while in female ovaries only a few molecular species were actively synthesized.     

Pollination of Nymphaea lotus (Nymphaeaceae) by Rhinoceros Beetles and Bees in the Northeastern Ivory Coast

Abstract: Detailed observations of the pollination biology of Nymphaea lotus were conducted by studying populations growing in seasonal ponds in the Comoé National Park (NE Ivory Coast). Flowers of N. lotus are protogynous and anthesis lasts for 4 to 5 days. Flowers open at sunset and close in the morning hours, thereby revealing a high variability in timing. For the first time, heat production in flowers could be observed in the genus Nymphaea. Flowers of N. lotus were visited by Ruteloryctes morio, an endemic dynastid beetle during the night. Beetles copulated in the flowers and fed on flower parts, but were less effective pollinators than several bee species that visited flowers in the early morning. Nymphaea lotus thus seems to be adapted to pollination by both nocturnal beetles and diurnal bees.     

The flower biology of Daphne gnidium L. (Thymelaeaceae)

Daphne gnidium is a circum-Mediterranean evergreen shrub. Flower and pollen features related to pollination mechanism were analyzed, and the seasonal presence of insects visiting the plant was monitored. The morphology and functionality of flowers are compatible with entomophily, as are pollen viability, which is maintained for a few hours only after anthesis, and the very short stigmatic receptivity. The reproductive fitness of the species is low and the ripening process often fails (35% collapsed embryos). While male organ functionality is fairly regular, the female organ often shows developmental arrest. The complex or transient evolutionary traits displayed by D. gnidium are comparable to those of Thymelaea hirsuta and other members of the Thymelaeaceae.     

Morphological diversity and function of the stigma in Ficus species (Moraceae)

The stigma plays several roles such as pollen hydration and selection, and pollen tube nutrition. In the Ficus-fig wasp mutualism, stigmata have an additional, almost unknown, function by representing a physical interface for both plant and wasp reproduction. We used light and electron microscopy to compare the detailed morphology of the stigmata of nine Ficus species of different sections and with different pollination modes and sexual expressions. Figs were collected at the stage when the stigmata were receptive for pollination. Stigmata in actively pollinated monoecious species have well developed papillae concentrated on the adaxial surface exposed towards the fig cavity. Conversely, the passively pollinated monoecious species have the whole surface of the stigmata covered by somewhat smaller papillae. In both actively and passively pollinated monoecious species these features are consistent, irrespective of style length. In all actively pollinated gynodioecious species, the stigmata of pistillate flowers were tubular or infundibuliform whereas in almost all actively pollinated monoecious species (except F. racemosa) the stigmata were filiform, with one branch or two asymmetric branches. In gynodioecious species the short-styled flowers in “male” figs show a limited receptive surface with small papillae, while the stigmata of long-styled flowers in “female” figs are covered by papillae that extend down the sides of the style, increasing the stigmatic surface. In actively pollinated species, stigmata are cohesive, forming a common surface for pollen tube germination (= synstigma). The synstigma arrangement was quite variable: lax, cohesive or very cohesive, with entanglement by stigmatic papillae and stylar trichomes. Entanglement by stylar trichomes is common in gynodioecious species. The synstigma arrangement did not correlate with phylogeny or breeding system. This study is the first to report a very loose synstigma in actively pollinated monoecious Ficus species. Our analyses revealed that, in Ficus, the synstigma is functionally analogous to an extra-gynoecial compitum. Comparative studies will be required to test further hypotheses about the evolutionary determinants of such variation.     

A Matter of Contrast: Yellow Flower Colour Constrains Style Length in Crocus species

Most flowers display distinct colour patterns comprising two different areas. The peripheral large-area component of floral colour patterns attracts flower visitors from some distance and the central small-area component guides flower visitors towards landing sites. Whereas the peripheral colour is largely variable among species, the central colour, produced mostly by anthers and pollen or pollen mimicking floral guides, is predominantly yellow and UV-absorbing. This holds also for yellow flowers that regularly display a UV bull’s eye pattern. Here we show that yellow-flowering Crocus species are a noticeable exception, since yellow-flowering Crocus species–being entirely UV-absorbing–exhibit low colour contrast between yellow reproductive organs and yellow tepals. The elongated yellow or orange-yellow style of Crocus flowers is a stamen-mimicking structure promoting cross-pollination by facilitating flower visitors’ contact with the apical stigma before the flower visitors are touching the anthers. Since Crocus species possess either yellow, violet or white tepals, the colour contrast between the stamen-mimicking style and the tepals varies among species. In this study comprising 106 Crocus species, it was tested whether the style length of Crocus flowers is dependent on the corolla colour. The results show that members of the genus Crocus with yellow tepals have evolved independently up to twelve times in the genus Crocus and that yellow-flowering Crocus species possess shorter styles as compared to violet- and white-flowering ones. The manipulation of flower visitors by anther-mimicking elongated styles in Crocus flowers is discussed.     

<Emphasis Type="Italic">Grias purpuripetala</Emphasis> (Lecythidaceae), a new purple-flowered species from southern Colombia

A new species, Grias purpuripetala, is described. It is the only species of the genus known to have the combination of monocaulous growth form; short inflorescence rachises; purple, erect petals at anthesis; anthers with small thecae and slightly apiculate connectives; and an intrastaminal disc. It is described based on collections from the western slopes of the Andes in the state of Nari?o in southern Colombia. The authors hypothesize that this spectacular species is bat- or beetle-pollinated because the flowers are nocturnal. This is the only species of the genus with purple petals―all others are either white or yellow.     

Structure of the stigma and style of <Emphasis Type="Italic">Callaeum psilophyllum</Emphasis> (Malpighiaceae) and its relation with potential pollinators

The family Malpighiaceae, particularly in the Neotropic, shows a similar floral morphology. Although floral attraction and rewards to pollinators are alike, stigmas and styles show more diversity. The stigmas were described covered with a thin and impermeable cuticle that needs to be ruptured by the mechanical action of the pollinators. However, this characteristic was only mentioned for a few species and the anatomy and ultrastructure of the stigmas were not explored. In this work, we analyze the morphology, anatomy, and ultrastructure of the stigma and style of Callaeum psilophyllum. Moreover, we identify the potential pollinators in order to evaluate how the disposition of the stigmas is related with their size and its role in the exposure of the receptive stigmatic surface. Our observations indicate that Centris flavifrons, C. fuscata, C. tarsata, and C. trigonoides are probably efficient pollinators of C. psilophyllum. The three stigmas are covered by a cuticle that remained intact in bagged flowers. The flowers exposed to visitors show the cuticle broken, more secretion in the intercellular spaces between sub-stigmatic cells and abundant electron-dense components inside vacuoles in stigmatic papillae. This indicates that the stigmas prepares in similar ways to receive pollen grains, but the pollinator action is required to break the cuticle, and once pollen tubes start growing, stigmatic and sub-stigmatic cells release more secretion by a granulocrine process.     

Inaccessible Biodiversity on Limestone Cliffs: Aster tianmenshanensis (Asteraceae), a New Critically Endangered Species from China

Aster tianmenshanensis G. J. Zhang & T. G. Gao, a new species of Asteraceae from southern China is described and illustrated based on evidence from morphology, micromorphology and molecular phylogeny. The new species is superficially similar to Aster salwinensis Onno in having rosettes of spatulate leaves and a solitary, terminal capitulum, but it differs by its glabrous leaf margins, unequal disc floret lobes and 1-seriate pappus. The molecular phylogenetic analysis, based on nuclear sequences ITS, ETS and chloroplast sequence trnL-F, showed that the new species was nested within the genus Aster and formed a well supported clade with Aster verticillatus (Reinw.) Brouillet et al. The new species differs from the latter in having unbranched stems, much larger capitula, unequal disc floret lobes, beakless achenes and persistent pappus. In particular, A. tianmenshanensis has very short stigmatic lines, only ca. 0.18 mm long and less than 1/3 of the length of sterile style tip appendages, remarkably different from its congeners. This type of stigmatic line, as far as we know, has not been found in any other species of Aster. The very short stigmatic lines plus the unequal disc floret lobes imply that the new species may have a very specialized pollination system, which may be a consequence of habitat specialization. The new species grows only on the limestone cliffs of Mt. Tianmen, Hunan Province, at the elevation of 1400 m. It could only be accessed when a plank walkway was built across the cliffs for tourists. As it is known only from an area estimated at less than 10 km² and a walkway passes through this location, its habitat could be easily disturbed. This species should best be treated as Critically Endangered based on the International Union for Conservation of Nature Red List Categories and Criteria B2a.     

Cleistogamy in the rare high Andean perennial herb Cryptantha capituliflora (Boraginaceae)

 Axial and apical flowers of Cryptantha capituliflora were analyzed with regard to morphology and pollen tube growth to assess the occurrence of cleistogamy. Although intermediate floral forms do occur, cleistogamous flowers were significantly smaller than chasmogamous flowers, had fewer anthers, and showed a distinctive stigmatic surface. Chasmogamous flowers can be cross-pollinated. Nevertheless, the growth of self-pollen tubes in few chasmogamous buds jointly with flower characters suggests that these flowers can probably produce fruits through autonomous selfing. The mean seed number per fruit did not differ between fruits from chasmogamous and cleistogamous flowers. Cleistogamous flowers were only observed in axial inflorescences, which are completely covered by the leaf. Other species of section Cryptantha also show the same trend, with cleistogamous flowers located in the lower half of the stems. This pattern is discussed in relation to dissimilarities in the outcrossing opportunities between flower types within the plant. Received May 22, 2002; accepted November 14, 2002 Published online: March 20, 2003     

Untersuchungen zur Blütenaffinität der Schachbrettweibchen (Agapetes galathea L.)

Rhopaloceran butterflies are well known to visit distinct species of flowers in order to extract nectar, normally leaving again after a short time to continue their foraging flights. Females of Agapetes galathea, however, often show a different behaviour, which is the subject of the following analysis. The dynamics of sugar production by flowers and the foraging pattern of the galathea- ♂♂ have been studied under natural conditions. Special consideration is given to the aggressive behaviour of the females towards approaching insects during their flower resting periods.     

Functional Significance of Labellum Pattern Variation in a Sexually Deceptive Orchid (Ophrys heldreichii): Evidence of Individual Signature Learning Effects

Mimicking female insects to attract male pollinators is an important strategy in sexually deceptive orchids of the genus Ophrys, and some species possess flowers with conspicuous labellum patterns. The function of the variation of the patterns remains unresolved, with suggestions that these enhance pollinator communication. We investigated the possible function of the labellum pattern in Ophrys heldreichii, an orchid species in which the conspicuous and complex labellum pattern contrasts with a dark background. The orchid is pollinated exclusively by males of the solitary bee, Eucera berlandi. Comparisons of labellum patterns revealed that patterns within inflorescences are more similar than those of other conspecific plants. Field observations showed that the males approach at a great speed and directly land on flowers, but after an unsuccessful copulation attempt, bees hover close and visually scan the labellum pattern for up to a minute. Learning experiments conducted with honeybees as an accessible model of bee vision demonstrated that labellum patterns of different plants can be reliably learnt; in contrast, patterns of flowers from the same inflorescence could not be discriminated. These results support the hypothesis that variable labellum patterns in O. heldreichii are involved in flower-pollinator communication which would likely help these plants to avoid geitonogamy.