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.     

Exine Micromorphology of Orchidinae (Orchidoideae,Orchidaceae): Phylogenetic Constraints or Ecological Influences?

BACKGROUND AND AIMS: Pollen characters have been widely used in defining evolutionary trends in orchids. In recent years, information on pollination biology and phylogenetic patterns within Orchidinae has become available. Hence, the aim of the presented work is to re-evaluate exine micromorphology of Orchidinae in light of recent phylogenetic studies and to test whether pollen micromorphology strictly depends on phylogenetic relationships among species or whether it is influenced by the marked differences in pollination ecology also reported among closely related species. METHODS: Pollen sculpturing of 45 species of Orchidinae and related taxa was investigated using scanning electron microscopy. To cover potential intraspecific variation, several accessions of the same species were examined. KEY RESULTS: Orchidinae show remarkable variation in exine sculpturing, with a different level of variation within species groups. In some genera, such as Serapias (rugulate) and Ophrys (psilate to verrucate), intrageneric uniformity corresponds well to a common pollination strategy and close relationships among species. However, little exine variability (psilate-scabrate and scabrate-rugulate) was also found in the genus Anacamptis in spite of striking differences in floral architecture and pollination strategies. A larger variety of exine conditions was found in genera Dactylorhiza (psilate, psilate-scabrate and reticulate) and Orchis s.s. (psilate, reticulate, perforate-rugulate and baculate) where no unequivocal correspondence can be found to either phylogenetic patterns or pollination strategies. CONCLUSIONS: Changes in pollen characteristics do not consistently reflect shifts in pollination strategy. A unique trend of exine evolution within Orchidinae is difficult to trace. However, the clades comprising Anacamptis, Neotinea, Ophrys and Serapias show psilate to rugulate or scabrate pollen, while that of the clade comprising Chamorchis, Dactylorhiza, Gymnadenia, Orchis s.s., Platanthera, Pseudorchis and Traunsteinera ranges from psilate to reticulate. Comparison of the data with exine micromorphology from members of the tribe Orchidieae and related tribes suggests a possible general trend from reticulate to psilate.     

Enantiostyly in Chamaecrista ramosa (Fabaceae‐Caesalpinioideae): floral morphology,pollen transfer dynamics and breeding system

Enantiostyly is a form of reciprocal herkogamy, in which floral morphs present reciprocal differences in the position of sexual elements, and occurs in monomorphic and dimorphic forms. This polymorphism maximises cross‐pollination and reduces self‐pollination, being very common within the subtribe Cassiinae (Fabaceae). Nevertheless, few studies have investigated the functionality of enantiostyly, particularly in this plant group. The present study aimed to investigate enantiostyly and its functionality in Chamaecrista ramosa, a monomorphic enantiostylous shrub, in an area of coastal vegetation in northeast Brazil. Pollen deposition and capture on the body of floral visitors, the relationship of these data with floral biology and breeding system, and morph ratio were evaluated. Pollen deposition and capture occurred in specific sites of the floral visitor body, showing the functionality of enantiostyly. The floral architecture, associated with the floral visitor behaviour, resulted in indirect pollen deposition on the floral visitor body. This occurred through a loop made by the pollen upon the inner petal surface, similar that generally reported for other Cassiinae. Chamaecrista ramosa is self‐compatible, although no fruit set was observed through spontaneous self‐pollination. The occurrence and number of floral morphs was similar within clumps. Enantiostyly seems to be advantageous for this species, as it results in efficient pollen capture and deposition, reduces the chances of autogamy and maximises intermorph pollen flow.     

Deconstructing a floral phenotype: do pollinators select for corolla integration in Lavandula latifolia?

An experimental field study was undertaken to assess whether correlational selection generated by pollinators can account for observed phenotypic integration of the two corolla lips in the zygomorphic flowers of the insect‐pollinated Mediterranean shrub, Lavandula latifolia (Labiatae). The experiment consisted of manipulating the upper and lower corolla lips according to a two‐way factorial design with interaction, and involved the complete (lower lip) or extensive (distal two thirds of upper lip) excision of one or the two lips. Per‐flower male (proportion of pollen grains removed) and female (number of pollen tubes down the style) pollination success, and maternal fecundity (seed set), of experimental flowers were evaluated after exposure to natural pollination in the field. Artificially induced corolla variation had no significant effects on either male or female pollination success, and two aberrant floral morphologies, characterized by lack of corolla integration, had a maternal fecundity advantage over normal two‐lipped phenotypes. These results do not support a predominant role of correlational selection by pollinators in maintaining corolla integration in L. latifolia, but are instead consistent with the alternative view that integration is mainly the consequence of genetic and/or developmental factors.     

Heterostyly and gene-flow in Menyanthes trifoliata L. (Menyanthaceae)

NIC LUGHADHA, E. M. & PARNELL, J. A. N., 1989. Heterostyly and gene-flow in Menyanthes trifoliata L. (Menyanthaceae). Menyanthes trifoliata is shown to be a self-incompatible heterostylous species with a suite of associated pollen and gynoecium dimorphisms. These features may promote legitimate pollination. Analysis of a natural population showed substantial pollen inviability, a preponderance of illegitimate pollen on stigmas and a possible asymmetry of pollen flow. Mean pollen travel was estimated as 2.6 m by counting legitimate grains on stigmas whose distance from the nearest source of legitimate pollen was known. Direct field and laboratory measurement indicated mean seed travel of <0.2m. The density of flowering spikes varied greatly within the studied colony of Menyanthes , with a mean of 0.93 m^-2. Using suitable corrections the genetically effective density was estimated as between 0.04 and 4.4 m^-2. Crossing experiments indicated that Menyanthes is fully outcrossed (r=1). The neighbourhood area was estimated as 10–15 m² while neighbourhood size ranged from less than one to sixty-five. Some published neighbourhood estimates are revised with the results for Menyanthes being at the lower end of the range. If the genetically effective density were estimated more accurately, neighbourhood estimates would likely be lower still. It is concluded that what is generally referred to as a population of Menyanthes , may actually represent many evolutionary units.     

The significance of flower colour change in eight co-occurring shrub species

Flower colour changes from white or yellow to various shades of red at or near the sites of harvestable pollen in Calytrix glutinosa, Grevillea pilulifera, Isopogon dubius and Petrophile biloba , and over most of the flower in Hypocalymma angustifolium, Verticordia chrysantha and V. huegelii and over the pseudanthium in Darwinia citriodora. All bee, wasp, beetle, fly, butterfly and moth visitors select flowers in the white/yellow phase rather than the red or intermediate phase.
Nectar is produced by five species, harvestable pollen by four species and detectable perfume by three species, all of which features are usually absent from the red phase. The timing of the colour change in all species also corresponds to loss of stigma receptivity, completion of pollination and onset of ovule seed) swelling. Six species also undergo minor morphometric changes which discourage visitation. In all species, colour change is non-inducible by pollinators, taking 2–30 days to complete. In three protandrous species, all available pollen may be removed in the first visit, requiring transport of non-self pollen to rewardless flowers during the 10 h period of the yellow phase.
These species are highly floriferous and occur in dense patches. Since only a small proportion of flowers may be receptive at any one time, it is concluded that retention of flower parts essentially serves to enhance long-distance attraction, while colour change maximizes pollination and foraging efficiency.     

Crane flies and microlepidoptera also function as pollinators in Epidendrum (Orchidaceae: Laeliinae): the reproductive biology of E. avicula

Crane flies and microlepidoptera have been recorded as pollinators in unrelated orchid groups, but these insects have never been recorded in Epidendroideae, the most species‐rich orchid subfamily, which includes one of the most diverse genera among Orchidaceae, Epidendrum. Based on data on phenology, floral morpho‐anatomy, pollinators, pollination mechanisms and breeding system, the reproductive biology of E. avicula was studied in south‐eastern Brazil. Epidendrum avicula possess osmophores that produce a citric fragrance at night. The flowers attract Tipulidae flies and several families of microlepidoptera that drink the nectar produced in a tube formed by the adnation of the labellum and column. As is common in Epidendrum, after removing the pollinarium, both crane flies and micro‐moths get trapped by the proboscis, which frightens the insects and inhibits any possible intent to immediately visit another flower. The behavior of the pollinators on flowers, plus the retention of the anther cap by the pollinarium, results in a reduction in the occurrence of geitonogamy. Because E. avicula is self‐incompatible, the consequence of pollinator behavior and the floral mechanisms tend to reduce the pollen loss. As far as we know, this is the first study to report the reproductive biology of a species of Epidendroideae pollinated by crane flies and microlepidoptera. Based on more recent concepts of plant–pollinator interactions, although E. avicula is pollinated by several species belonging to two distinct orders, suggesting an unspecialized pollination system is involved, nectar‐seeking microlepidoptera and Tipulidae flies can be recognized as a single functional group.     

Pollination in conifers

Our understanding of pollination in conifers has advanced rapidly in recent years, but it still lags behind our knowledge of this process in angiosperms. In part this is because conifers are not considered to be high priority crops and, unlike many cultivated flowers, conifer seed cones are generally neither large nor colorful. The use of genetics to improve tree growth has primarily been through selection and asexual propagation rather than breeding, and because incompatibility is not thought to occur in conifer pollination systems, concern about pollination has primarily been with regard to seed production. Here we examine the ancestral wind-pollination mechanism in conifers and discuss how the process may have evolved to improve pollination success.     

Pollen-size comparisons among animal-pollinated angiosperms with different pollination characteristics

Pollen volume varies among angiosperm species over five orders of magnitude, presumably because the functional advantages of pollen size depend on each species' particular pollination and fertilization conditions. This paper reports two studies that assess whether animal-pollinated species with different pollination systems differ correspondingly in pollen size, as would be expected if pollen size affected pollen transport. Analysis of nine Pedicularis species detected significant interspecific variation in pollen size; however, the overall mean pollen volume of species pollinated primarily by nectar-collecting bees did not differ significantly from that of species pollinated by pollen-collecting bees. Similarly, comparison of bee- and bird-pollinated congeners in 16 genera (nine families) from Australia and North America found considerable variation in pollen diameter within and between genera, but this variation was not associated consistency with differences in primary pollinator type (bee>bird for three genera, bee > bird for five genera, bee相似文献   

Pollination biology of the endangered orchid Cypripedium japonicum in a fragmented forest of Japan

Pollination biology studies of the endangered orchid Cypripedium japonicum were conducted in its natural habitat using pollinator observation and hand‐pollination experiments. The observed fruit set was as follows: artificial outcross‐pollinated, 100%; artificial self‐pollinated, 100%; pollinator‐excluded, 0%; and emasculated flowers, 0%. These results show that this species, although self‐compatible, is neither autogamous nor agamospermous. The fruit set for open‐pollinated flowers was 14.9%, which suggests that the study population was subject to pollinator limitation. The nectarless flowers of C. japonicum were exclusively visited and pollinated by the queens of two bumblebee species (Bombus ardens and B. diversus diversus). It is probable that the nectarless flowers of C. japonicum attract pollinators through a generalized food deceptive system.     

毛菍传粉生物学研究

为探讨毛菍(Melastoma sanguineum Sims.)传粉生物学特性,对海南霸王岭国家级自然保护区林缘、林内和林隙3个生境内毛菍的花期、花部形态、访花昆虫等进行观察,同时对其繁育系统相关指标进行了测定。结果表明：毛菍无花蜜分泌;花瓣对访花昆虫有吸引作用;访花昆虫主要有蜂类、食蚜蝇类和丽金龟类;主要传粉昆虫有木蜂、熊蜂,其中木蜂传粉活动最为活跃,为最有效传粉昆虫。毛菍具典型的异型雄蕊,长、短雄蕊在传粉过程中具有明确的功能分化,推测这是毛菍保证传粉成功的策略。毛菍为自交亲和的异交种,需要昆虫传粉;人工自交和异交均具有较高的坐果率;不存在无融合生殖、主动自花授粉和滞后自交的生殖保障现象;其繁育系统是兼性自交。昼夜温差较小和相对湿度较高的林内和林隙生境有利于延长群体花期;光照充足、温度相对较高的林缘生境,花朵发育较快,柱头可授性的时间稍早。花粉、传粉者和环境条件是制约毛菍野外传粉效率的主要因素。     

巴旦杏授粉试验及花粉管生长的荧光显微观察

对巴旦杏4个主栽品种的授粉试验结果表明：自花授粉试验中“鹰嘴”、“双果”坐果率分别为19．74％和4．29％,“纸皮”和“麻壳”坐果率为零;自然授粉状态下4个主栽品种坐果率在9．37％－20．13％之间;人工异花授粉试验中4个品种9个授粉组合坐果率绝大多数都在40％以上,其中混合花粉授粉试验结果最好;人工养蜂条件下的坐果率明显高于无蜂自然授粉的坐果率,昆虫传粉可达到人工混合花粉授粉的效果。授粉试验还表现出巴旦杏雌蕊柱头的有效授粉时间为开花后花期的前4d,授粉越早,越有利于坐果。荧光显微观察表明,巴旦杏花粉管最早约于授粉第7天通过花柱基部到达子房室,第8天前后到达胚珠。     

Heterostyly promotes compatible pollination in buckwheats: Comparisons of intraflower,intraplant, and interplant pollen flow in distylous and homostylous Fagopyrum

Premise of the Study

Heterostyly, the reciprocal positioning of stigmas and anthers in different floral morphs, has long been thought to promote intermorph pollination. However, extensive intramorph pollination occurs commonly in heterostylous species, leading to recurrent questions about the functional and evolutionary significance of heterostyly.

Methods

To identify the sources of stigmatic pollen (autogamous [intraflower], geitonogamous [intraplant], vs. interplant), we emasculated either one flower or entire plants in experimental populations of the two closely related buckwheat species, distylous Fagopyrum esculentum and homostylous F. tataricum. Differences in pollen size allowed unambiguous identification of pollen on stigmas.

Results

Only 2.4% of F. tataricum pollen and 1.5% of F. esculentum pollen arrived successfully on compatible stigmas of other plants. In the former (homostylous) species, 71.3% of the pollen load on stigmas was autogamous, 10.8% was geitonogamous, and 17.9% was interplant. In the latter (distylous) species, 37.45% of the pollen on stigmas was autogamous, 13.8% was geitonogamous, 17.0% was intramorph, and 31.75% was intermorph. The amount of incompatible pollen arriving on stigmas was greatly decreased by both one‐flower and whole‐plant emasculations, and thus, the proportion of compatible pollen deposited increased with one‐flower emasculation and increased even more with whole‐plant emasculation.

Conclusions

Our quantification of pollen‐donor sources in these two species indicated that heterostyly in Fagopyrum esculentum provided a nearly 2‐fold fitness advantage (in terms of compatible pollination) over expected (random) pollen transfers between morphs. Because of reduced herkogamy, the homostylous F. tataricum was highly autogamous.     

Types of pollen dispersal units in orchids,and their consequences for germination and fertilization

The various pollen dispersal units (PDU) found in orchids are discussed together with possible evolutionary trends and the consequences for germination and fertilization. Orchids with monad and tetrad pollen form more complex dispersal units by means of pollenkitt, elastoviscin, a callosic wall, common walls or a combination of these. Evolutionary trends include (1) from pollenkitt to elastoviscin; (2) from monad to tetrads and multiples of tetrads; (3) from partially dehydrated (<30 %) to partially hydrated (>30 %) pollen; and (4) from monad pollen to PDUs with many pollen grains. The biological consequences concern both male and female reproductive systems. Some features of the male side are present in all orchids irrespective of the pollen dispersal unit, whereas other characters are found only in orchids with pollinia; the same applies for the female counterpart. Pollen grains of orchids with pollinia germinate at least 24 h after pollination because the pollen grains/tetrads must swell and make space for the growth of pollen tubes.     

国家二级保护植物翅果油树传粉生物学的初步研究

对翅果油树进行了传粉生物学的野外观察和实验表明：（1）翅果油树的花具有自花传粉的结构特征,同时具有蜜腺,又具备异花传粉的特征。传粉者主要是一种家养的蜜蜂。（2）翅果油树花形态结构与传粉者的形态和传粉行为非常协调和适应,昆虫的传粉部位主要是头前部和胸部。（3）蜜腺的产蜜量大,开花后1～2天蜜腺分泌量最大。对传粉昆虫竞争中具明显优势。昆虫的访花频率明显高于同花期的其他植物。（4）翅果油树没有融合生殖现象。