POLLINATION OF RAFFLESIA (RAFFLESIACEAE)

The genus Rqfflesia includes about 13 species of parasitic flowering plants, among which are the largest known flowers. The flower with subtending scales is the only part of the plant external to the host and is produced solitary on roots (rarely stems) of the genus Tetrastigma (Vitaceae). Field studies were made of the pollination process in R. pricei, a species endemic to the Crocker Range in the Malaysian state of Sabah (northern Borneo). Pollination is mediated by carrion (bluebottle) flies of the genera Lucilla and Chrysomya. Experimental data indicate that both visual and olfactory cues are important in attracting flies to flowers. Flies (mostly female L. papuensis) obtain loads of the viscous liquid pollen matrix by visiting male flowers and entering anther grooves on the central column of the flower, precisely guided by ridges armed with hairs that force the fly into a position in which the pollen is positioned on the dorsal part of the thorax. “Windows” on the inside of the perigone diaphragm apparently help orient their flight inside flowers. Pollen-loaded flies visiting female flowers may enter the infradiscoidal sulcus formed by a broad ring of stigmatic tissue above and the expanded base of the column below. On entering the sulcus the fly is wedged in so tightly that pollen is rubbed off the thorax onto the stigma. Only large flies could be effective in picking up pollen from male flowers and transferring it to female flowers. The pollination syndrome is sapromyophily, in which the flower closely parallels trap flowers of several other plant families, although it is not a trap. The flower provides no reward for pollinators but deceives them by an apparent offering of food and possibly brood place. Rafflesia plants are extremely rare, perhaps in part because of infrequency of pollination, which requires neighboring male and female flowers simultaneously in bloom.     

Insect pollination and floral mechanisms in South African species ofSatyrium (Orchidaceae)

The morphologically diverse flowers in the genusSatyrium reflect adaptations to a wide range of pollinators. Several recently discovered pollination systems inSatyrium are described and illustrated here; these include pollination by solitary bees, carrion flies, butterflies and moths. Two basic types of floral mechanism are recognised inSatyrium: (1) Species pollinated by lepidoptera and birds have long floral spurs and plate-like viscidia seated in lateral rostellum notches; these viscidia become attached to the proboscis or bill of the pollinator. (2) Species pollinated by flies and bees have relatively short floral spurs and globose viscidia seated in terminal rostellum notches; these viscidia become attached to the face, thorax or eyes of the pollinator.     

High temperatures applied at fall forcing disturb ovule development in Syringa vulgaris L. “Mme Florent Stepman”

Under natural conditions, Syringa vulgaris blooms in May but it may be forced to produce blooming panicles in autumn, winter and early spring. To break winter dormancy, plants require high temperatures, the range of which depends on the forcing date. Forcing in November requires exposure to temperatures of ∼37°C. Such an exposure may affect the development of both male and female flower organs. In this work we have followed the anatomical changes in the ovule development in buds collected from plants forced in autumn, and compared them to those in plants blooming under natural conditions in May. From the earliest identifiable stages, in flowers forced in November the ovule development was delayed relative to the normal development pattern. Ultimately, most of the ovules (83%) degenerated before the differentiation of the embryo sacs was completed. We believe that it is the high temperatures required during autumn forcing that affect the development of female gametophytes, leading to the formation of defective structures which often degenerate. This in turn may reduce the esthetic appeal of the inflorescences and reduce their commercial value.     

Carrion mimicry in a South African orchid: flowers attract a narrow subset of the fly assemblage on animal carcasses

BACKGROUND AND AIMS: Although pollination of plants that attract flies by resembling their carrion brood and food sites has been reported in several angiosperm families, there has been very little work done on the level of specificity in carrion mimicry systems and the importance of plant cues in mediating such specialization. Specificity may be expected, as carrion-frequenting flies often exploit different niches, which has been interpreted as avoidance of interspecific competition. Interactions between the orchid Satyrium pumilum and a local assemblage of carrion flies were investigated, and the functional significance of floral traits, especially scent, tested. Pollination success and the incidence of pollinator-mediated self-pollination were measured and these were compared with values for orchids with sexual- and food-deceptive pollination systems. METHODS AND KEY RESULTS: Observations of insect visitation to animal carcasses and to flowers showed that the local assemblage of carrion flies was dominated by blow flies (Calliphoridae), house flies (Muscidae) and flesh flies (Sarcophagidae), but flowers of the orchid were pollinated exclusively by flesh flies, with a strong bias towards females that sometimes deposited live larvae on flowers. A trend towards similar partitioning of fly taxa was found in an experiment that tested the effect of large versus small carrion quantities on fly attraction. GC-MS analysis showed that floral scent is dominated by oligosulfides, 2-heptanone, p-cresol and indole, compounds that also dominate carrion scent. Flesh flies did not distinguish between floral and carrion scent in a choice experiment using olfactory cues only, which also showed that scent alone is responsible for fly attraction. Pollination success was relatively high (31·5 % of flowers), but tracking of stained pollinia also revealed that a relatively high percentage (46 %) of pollen deposited on stigmas originates from the same plant. CONCLUSIONS: Satyrium pumilum selectively attracts flesh flies, probably because its relatively weak scent resembles that of the small carrion on which these flies predominate. In this way, the plants exploit a specific subset of the insect assemblage associated with carrion. Pollination rates and levels of self-pollination were high compared with those in other deceptive orchids and it is therefore unlikely that this mimicry system evolved to promote outcrossing.     

Correction to: The floral biology and the role of staminal connective appendages during pollination of the endoparasite Bdallophytum americanum (Cytinaceae)

Bdallophytum americanum (Cytinaceae) is an endoparasitic plant species, meaning only the flowers emerge from the host during the reproductive season. Reports on the pollination biology of this species state that its primary pollinators are carrion flies attracted by the smell of the flowers and nectar as a reward. However, the functional role of one of the most outstanding attributes of B. americanum has been neglected. These are the staminal appendages formed by the apical overgrowth of connective tissue during anther development. To determine whether these staminal appendages play a role in pollination, we monitored a nectarless population of B. americanum. We described the inflorescence emergence, floral movements, and pollination and performed field experiments to test whether the absence of the staminal connective appendages affected the visitation frequency. Male inflorescences emerge early, and both male and female flowers open during the day and do not close. Hoverflies are the most frequent visitors to both floral sexes and carry the most pollen. Moreover, the movement of staminal appendages matching the pollen viability changes is reported for the first time. The staminal appendages are the structures where pollinators land before foraging. The field experiments showed that the visitation frequency decreased sharply without staminal appendages. As a landing platform, the staminal connective appendages in B. americanum are crucial for pollinator positioning and collecting viable pollen.

     

Dioecy inHonkenya peploides var.major (Caryophyllaceae)

Sexual difference and pollination system inHonkenya peploides var.major were investigated. The present analyses indicate that sex expression of this plant is functionally dioecious, rather than androdioecious, as described previously. Its flowers are largely pollinated by small bees, hover-flies, flies, and ants. Female flowers were morphologically characterized by conspicuously longer styles and smaller petals, and had no pollen grains, while male flowers had shorter styles, larger petals, and longer filaments, and produced viable pollen grains. Although the male flowers possess many ovules in an ovary, they set fruits at a very low frequency in natural conditions and did not produce fruits by pollination experiments. We suggest that the dioecy of this species has been derived from hermaphrodite through gynodioecy, and not directly from hermaphrodite.     

The pollination of the gynomonoecious Bdallophytum oxylepis (Cytinaceae,Malvales)

Bdallophytum oxylepis is a rare and endemic species belonging to the Cytinaceae family, a root holoparasitic plant in which most resources are allocated to attracting pollinators. This species is gynomonoecious with intraindividual variation in flower size and sex. Moreover, the flowers exhibit sapromyophilous traits, as do other species of Bdallophytum. Firstly, this study aimed to determine whether all floral morphs can form seeds and be pollen donors (in the case of bisexual flowers). Secondly, as this species has floral traits hypothesized to adapt to particular types of pollen vectors (carrion flies), we also studied the pollination of B. oxylepis to confirm whether the syndromes correspond to what occurs in nature. Through pollination treatments, we determined that all floral morphs are functional. By monitoring the inflorescences, we found that pollination is specialized in the studied population. Stingless bees performed pollination, as they have a high visitation rate, frequency, and constancy, and they are unique visitors that deposit pollen on the stigmas. Thus, they appear to be effective pollinators rather than carrion flies, as predicted by the syndrome. As shown here, animal–plant interaction studies can help establish a basis for conserving rare species such as holoparasites. Moreover, knowledge about the reproductive aspects of B. oxylepis reveals essential clues about its life cycle and role in maintaining native pollinators with economic and cultural value, such as stingless bees.     

Convergent evolution of carrion and faecal scent mimicry in fly-pollinated angiosperm flowers and a stinkhorn fungus

Flowers of many angiosperms attract fly pollinators through mimicry of animal carrion and faeces. This phenomenon of “sapromyiophily” is also evident in the sporophytes of some mosses and fruiting bodies of “stinkhorn” fungi, both of which use flies as agents of spore dispersal. We studied the scent chemistry of a stinkhorn fungus (Clathrus archeri) and seven fly-pollinated plant species with foetid odours to determine the degree to which these organisms mimic the scent of carrion and faeces (reference scent samples were collected from rotting meat, a rat carcass and horse and dog faeces), as well as the degree of convergent evolution between the fungus and angiosperm flowers. We found that scents of both the fungus and angiosperms tended to contain compounds typical of carrion, such as oligosulphides, and of faeces, such as phenol, indole and p-cresol. This study provides compelling new evidence for mimicry of carrion and faeces, as well as a striking pattern of convergence in the putrid scents of the fungus and the angiosperms, relative to those of confamilial species. The syndrome of sapromyiophily thus encompasses at least two kingdoms (Plantae and Fungi) and provides an effective means of exploiting flies as agents of pollen and spore dispersal.     

金沙江干热河谷滇榄仁开花物候与繁育系统

通过野外定点观测,并利用套袋实验、解剖实验以及花粉形态特征观察与活力检测、柱头可受性检测、花粉/胚珠比以及异交指数等,研究了金沙江干热河谷滇榄仁自然群体花部形态特征与开花进程、繁育系统特征以及传粉媒介与环境的适应性等内容。结果表明,滇榄仁始花期在每年4月上旬,群体花期持续时间为30d左右,开花同步性高,呈典型的"集中开花模式"。单花开放时,柱头先伸长和膨大,约4d后雄蕊伸出,具有雌雄异熟的特征,一定程度上避免了自交的发生。单花开放进程可划分为花蕾期、柱头伸长期、雄蕊始露期、盛开期、盛开后期、花谢期等6个时期。除了花蕾期外,整个开花期的花粉均具有活力。花粉活力与柱头可受性之间避开了最适授粉期,但也有部分重叠。滇榄仁可能同时具有风媒和虫媒传粉。繁育系统以异交为主,部分自交亲和,需要传粉者。滇榄仁花部特征、开花物候和繁育系统为其适应干热河谷恶劣气候环境提供了一定的生殖保障和进化潜力。良好的群落环境有助于促进滇榄仁异花传粉,产生更多有效的种子,从而促进林下自然更新。     

Pollination of the “carrion flowers” of an African stapeliad (<Emphasis Type="Italic">Ceropegia mixta</Emphasis>: Apocynaceae): the importance of visual and scent traits for the attraction of flies

Flowers that mimic carrion or faeces exhibit unusual traits, the evolution and functional significance of which remain poorly understood. Odour is an important pollinator attractant, but visual traits and interactions between visual and scent traits have seldom been considered. We studied pollination of the “carrion flowers” of Ceropegia mixta [= Orbea variegata], analysed floral traits and used manipulative experiments to explore the contributions of visual and scent traits to pollinator attraction. Flowers were pollinated primarily by Musca domestica (Muscidae), with lesser contributions by Calliphoridae and Sarcophagidae flies. The floral odour (analysed using gas chromatography–mass spectrometry) was dominated by oligosulphides and phenol. Comparison of floral and abiotic background colours (analysed using reflectance spectrometry) using a fly colour vision model suggested that flowers would be chromatically indistinguishable from the background. Comparison of fly arrival rates at concealed (but still scented) versus exposed flowers showed that flies can locate flowers without visual cues, but visitation was higher when the flowers were visible. Experiments using model flowers with odour supplied by real flowers (to explore the significance of dark flowers and dark spots on a pale background, which both occur frequently in flowers that mimic carrion or faeces) showed that scented black flowers attracted significantly more flies than similarly scented human-yellow flowers, while the presence or size of black spots on the corolla had no effect on the attraction of flies. Our results suggest that there is a visual component to fly attraction, but some traits, such as the mottled patterning, may not have evolved to enhance pollinator attraction.     

Synchronized protandry and hermaphroditism in a tropical secondary forest tree, Schefflera heptaphylla (Araliaceae)

Selection favoring avoidance of stigma clogging, pollen discounting, self-fertilization, and other negative effects of self-pollination can produce intricate patterns of intra- and interfloral dichogamy in plants bearing numerous flowers. Here we report an extensive study of the relationships among dichogamy, floral sex allocation (pollen-to-ovule ratios), nectar production, floral visitors, mating system, and fruit set in natural populations of Schefflera heptaphylla, a widespread paleotropical secondary forest tree that produces thousands of flowers in a blooming season. Each tree produces 15?C30 sequentially blooming, paniculate, compound inflorescences. Each compound inflorescence has up to three orders of umbellets, which also bloom sequentially. While hand-pollinations showed that S. heptaphylla was capable of self-fertilization, our observations of thousands of flowers showed that strong intra- and interfloral protandry severely restricts both autogamous and geitonogamous self-pollination. All flowers were bisexual, thus the sexual system of the populations we studied was hermaphroditism. The pollen-to-ovule (P/O) ratios were characteristic of outcrossing species, and P/O ratios of flowers in the last-maturing (third order) umbellets were significantly higher than those in earlier-maturing (first and second order) umbellets. Floral visitors were primarily flies (Chrysomya sp. and Syrphinae sp.) and wasps (Vespula sp. and Eumenes sp.). Flowers produced nectar during both the male (pollen presentation) and female (stigma receptivity) stages of their development, and the volume of nectar production was higher in the female stage. Nevertheless, flowers received fewer visits in the female stage than they did in the male stage, and natural fruit set was low, especially in first and third order umbellets. Fruit set from hand cross- and self-pollinations was significantly higher than natural fruit set, indicating pollen limitation of fruit set. Schefflera heptaphylla has also been reported to be andromonoecious. Both hermaphroditism and andromonoecy are consistent with theoretical predictions for variation in sex allocation among sequentially maturing flowers in protandrous species. Further studies comparing hermaphroditic and andromonoecious populations of S. heptaphylla could elucidate the selective factors affecting sex expression, nectar production, and fruit set in species with numerous flowers displaying both intra- and interfloral dichogamy.     

Targeted detection of mammalian species using carrion fly‐derived DNA

DNA analysis from carrion flies (iDNA analysis) has recently been promoted as a powerful tool for cost‐ and time‐efficient monitoring of wildlife. While originally applied to identify any mammalian species present in an area, it should also allow for targeted detection of species and individuals. Using carrion flies captured in the Taï National Park, Côte d'Ivoire, we assessed this possibility by (i) screening carrion fly DNA extracts with nonspecific and species‐specific PCR systems, respectively, targeting mitochondrial DNA (mtDNA) fragments of any mammal or of Jentink's duiker (Cephalophus jentinki), three colobine monkeys (subfamily Colobinae) and sooty mangabey (Cercocebus atys); and (ii) genotyping carrion fly extracts containing sooty mangabey mtDNA. In comparison with the nonspecific PCR assay, the use of specific PCRs increased the frequency of detection of target species up to threefold. Detection rates partially reflected relative abundances of target species in the area. Amplification of seven microsatellite loci from carrion flies positive for sooty mangabey mtDNA yielded an average PCR success of 46%, showing that the identification of individuals is, to some extent, possible. Regression analysis of microsatellite PCR success and mtDNA concentration revealed that, among all carrion flies analysed for this study, 1% contained amounts of mammal mtDNA sufficient to attempt genotyping with potentially high success. We conclude that carrion fly‐derived DNA analysis represents a promising tool for targeted monitoring of mammals in their natural habitat.     

Insect pollinators show constancy for different flower traits between the most- and less-preferred plants: a case study of the long-proboscid tangle-veined fly

• The coevolution of insect pollinators and their host plants is a typical example of natural selection; however, how insect pollinators avoid overdependence on one peculiar plant remains unclear. As most insect pollinators exhibit a diet breadth when showing flower constancy, determining the difference and similarity of most and less preferred flowers by insect pollinators may be helpful to understand their trade-off between flower constancy and overdependence.
• This was addressed using the long-proboscid tangle-veined fly (Nemetrinus spp.). Dietary investigation indicates that the flies show constancy for the morphological characteristic of the Delphinium caeruleum, which is the most preferred plant for this Nemestrinidae fly that has blue, long-tubed flowers.
• In a colour selection experiment, focal individuals showed obvious preference for white, which is the colour of less preferred flowers by the fly in the natural environment. In a scent selection experiment, focal individuals showed obvious preference for D. caeruleum and Dracocephalum heterophyllum but avoidance to Dasiphora fruticosa and Dasiphora davurica. This indicates that long-proboscid tangle-veined flies can forage on other flowers, despite the existing constancy for D. caeruleum, as long as they do not hate the scent. It seems that long-proboscid tangle-veined flies can maximise foraging efficiency by showing constancy for the morphological characteristic of the most preferred plant and for the scent and colour of less-preferred plants.
• The trade-off of long-proboscid tangle-veined flies in the selection of nectar sources may be an adaptation to the risk of overdependence on one plant in evolution.

     

Pollen limitation in three sympatric species ofVaccinium (Ericaceae) in the Upper Ardennes, Belgium

The reproductive success of three co-flowering species ofVaccinium (V. myrtillus, V. vitis-idaea andV. uliginosum) was studied in one heathland of the Upper Ardennes, Belgium, during three years (1988–1990). The purpose was to examine whether pollen limitation, flower position and flowering phenology may influence patterns of fruit set in these three sympatric species. I quantified fruit and seed set following supplementary hand-pollinations and compared this to natural fruit set. On the same plants, I also quantified fruit and seed set in relation to the spatial position of the flowers on the ramet and their temporal sequence of blooming. Hand-pollination had no significant effect on fruit set inV. vitis-idaea andV. uliginosum, but significantly increased seed number per fruit in 1989. InV. myrtillus both fruit and seed set were increased by supplementary pollination, but significantly in only one year. Analyses of position effects revealed that the fruits in the lower positions in the ramet did not mature preferentially and did not contain more seeds inV. uliginosum and inV. vitis-idaea. Flowering phenology also had no significant effect for these species. InV. myrtillus hand-pollinated flowers showed a seasonal decline in seed number, and control (naturally pollinated) flowers showed a seasonal increase in fruit set. Fruit and seed set appear to be pollen-limited rather than resource-limited inV. uliginosum. In the two other species, unfavorable weather (frosts) could be a more important cause of low fruit and seed set.     

Thermal ecophysiology of seven carrion-feeding blowflies in Southern Africa

A variety of temperature thresholds for larvae, pupae, and adults of seven African species of carrion‐feeding blowflies (Diptera: Calliphoridae) was measured and compared to understand their basic thermal biology and the influence of temperature on their behaviour. Calliphora croceipalpis (Jaennicke) had consistently lower temperature thresholds than all other species tested for all larval (42.9 °C), pupal (16.6 °C), and adult (45.6 °C) stages. Larvae (50.1 °C) and adults (53.4 °C) of Chrysomya marginalis (Robineau‐Desvoidy) had higher upper lethal temperature thresholds than all other species and weighed more than all other species. Pupae and adults of both Chrysomya albiceps (Wiedemann) and Lucilia sericata (Meigen) had similar temperature thresholds, whereas Chrysomya putoria (Wiedemann), Chrysomya chloropyga (Wiedemann), and Chrysomya megacephala (Fabricius) had inconsistent rank temperature thresholds between the larval, pupal, and adult stages. With a few minor exceptions, the nervous activity, muscle activity, and death thresholds in female adult flies responded at higher temperatures than conspecific male flies for all species tested. Similarly, female adult flies weighed consistently more than conspecific male flies for all species tested, except Ca. croceipalpis. These data suggest that there is a phylogenetic component to the thermal biology of blowflies, because Ca. croceipalpis belongs to a primarily Holarctic genus and shows adaptation to that climate even though it inhabits Africa. Comparisons between these temperature thresholds and the distributions of blowfly species present on three rhinoceros carcasses suggest that blowfly larvae with high upper lethal temperature thresholds (particularly C. marginalis) dominate in interspecific competition on the carcass by raising the temperature of the amassed maggots above the thresholds of other carrion‐feeding blowflies, through metabolically generated heat.     

Bimodal cue complex signifies suitable oviposition sites to gravid females of the common green bottle fly

Gravid females of the common green bottle fly, Lucilia sericata Meigen (Diptera: Calliphoridae), readily locate recently deceased vertebrates as oviposition sites, particularly when these animals have been injured. We investigated semiochemical and visual cues that mediate attraction of gravid females to fresh rat carrion. Female flies were more strongly attracted to incised rat carrion than to intact carrion. They were also attracted to Porapak Q headspace volatile (HSV) extract of incised rat carrion. Analyzing aliquots of Porapak Q HSV extract by gas chromatographic‐electroantennographic detection revealed nine components [phenol, para‐ and/or meta‐cresol (could not be separated), guaiacol, dimethyl trisulfide (DMTS), phenylacetaldehyde, (E)‐2‐octenal, nonanal, and tetramethyl pyrazine] that consistently elicited responses from blow fly antennae. In laboratory experiments, a synthetic blend of these nine components was as attractive to gravid females as Porapak Q HSV extract, but blend attractiveness was due entirely to DMTS. In both laboratory and field experiments, increasing doses of DMTS attracted increasingly more flies. Coupled with DMTS, carrion‐type color cues (dark red, black) were more effective than bright color cues (white, yellow) in attracting flies. In field experiments, dark traps baited with DMTS captured a total of 214 calliphorid flies (200 L. sericata, 10 Lucilia illustris Meigen, three Calliphora vicina Robineau‐Desvoidy, one Calliphora vomitoria L.), all of which were gravid females. These results support the conclusion that DMTS and dark color represent a bimodal cue complex that signifies suitable oviposition sites to gravid calliphorid females, particularly L. sericata.     

POLLEN AND GENE DISPERSAL: THE INFLUENCES OF COMPETITION FOR POLLINATION

Pollinators that forage indiscriminately can transfer pollen from one species to another, reducing the amount that reaches conspecific flowers. I present evidence that the presence of another plant species visited by the same pollinators can also reduce pollen dispersal distances and outcrossing. This has the potential to influence gene flow and reproductive success. Pollen carryover and movement patterns were measured for the shared insect pollinators of Stellaria pubera and Claytonia virginica in North Carolina. Bee flies deposited similar amounts of Stellaria pollen on a series of pistillate Claytonia flowers as on a series of pistillate Stellaria flowers. In arrays of potted plants, flies and solitary bees visited most flowers on a plant before leaving and then flew to a nearby plant chosen independently of species; 95% of moves were to one of 12 nearest neighbors. These measures of pollen carryover and movement patterns were used in a set of computer simulations to predict pollen dispersal distances. The simulations suggested that C. virginica substantially reduces outcrossing and pollen flow in S. pubera. These predictions were tested by tracking dye movement from anthers in populations of potted plants. Addition of C. virginica reduced the mean squared distance moved by dye to receptive S. pubera flowers by 23% and reduced the amount of dye moved by 51%. The estimated pollen component of gene flow was also much lower in a natural population of 5. pubera mixed with C. virginica than in the synthetic single-species populations.     

Stage-specific localization of the small heat shock protein Hsp27 during oogenesis inDrosophila melanogaster

The developmental and heat shock-induced expression of the small heat shock protein Hsp27 was investigated by confocal microscopy of whole-mount immunostained preparations of ovarioles during oogenesis inDrosophila melanogaster. In unstressed flies, Hsp27 was mainly associated with germline nurse cells throughout egg development. A small group of somatic follicle cells also expressed Hsp27 specifically at stages 8 to 10 of oogenesis. Interestingly, this Hsp showed a different intracellular localization depending on the stages of egg chamber development. Thus Hsp27 was localized in the nucleus of nurse cells during the first stages of oogenesis (from germarium to stage 6) whereas it showed a perinuclear and cytoplasmic localization from stage 8. After a heat shock, Hsp27 accumulated in somatic follicle cells surrounding the egg chamber whereas the expression of this small Hsp did not seem to be enhanced in nurse cells. The stage-dependent pattern of intracellular localization of Hsp27 observed in nurse cells of unstressed flies was also observed following heat shock. At late stages of oogenesis, Hsp27 also showed a perinuclear distribution in follicle and nurse cells after heat stress. These observations suggest that different factors may modulate the expression and intracellular distribution of Hsp27. This modulation may be associated with the specific activities occurring in each particular cell type throughout oogenesis during both normal development and under heat shock conditions. Edited by: E.R. Schmidt     

Pollinator directionality as a response to nectar gradient: promoting outcrossing while avoiding geitonogamy

Plants with multiple flowers could be prone to autonomous self‐pollination and insect‐mediated geitonogamy, but physiological and ecological features have evolved preventing costs related to autogamy. We studied the rare perennial herb Dictamnus albus as a model plant, with the aim of describing the plant–pollinator system from both plant and pollinator perspectives and analysing features that promote outcrossing in an entomophilous species. The breeding system and reproductive success of D. albus were investigated in experimental and natural conditions, showing that it is potentially self‐compatible, but only intra‐inflorescence insect‐mediated selfing is possible. Nectar analysis showed gender‐biased production towards the female phase, which follows the male phase, and during flowering, full blooming is found in flowers at the bottom of the raceme. Among a wide spectrum of insect visitors, three genera (Bombus, Apis, Megachile) were found to be principal pollinators. A study of insect behaviour showed a tendency towards bottom‐to‐top flights for the most important pollinators Bombus spp. and Apis mellifera: upward movements on the racemes could be explained by foraging behaviour, from more to less rewarding flowers. In accordance with the ‘declining reward hypothesis’, bumblebees and honeybees leave the plant when gain of reward is low, after which few flowers are visited, reducing the chance of self‐pollen transfer among flowers. Intra‐flower self‐pollination is prevented in D. albus by protandry and herkogamy, while the nectar‐induced sequential pattern of pollinator visits avoids geitonogamy and tends to maximise pollen export, promoting outcrossing. All these features for preventing selfing benefit plant fitness and population genetic structure.     

Effect of Flowering Stages on the Content of Active Ingredients and Antioxidant Capability of Bletilla striata Flowers

Bletilla striata (Thunb.) Reichb.f. is a perennial herb with abundant active ingredients. Previous research mainly focused on its tubers, however, the study on flowers, especially the variation of active ingredient contents at different flowering stages, was rarely seen. This study analyzed the total phenols, flavonoids, polysaccharides, anthocyanins, and cyanidin-3-O-glucoside content of B. striata flowers which were in cultivated in Herb Garden of Zhejiang A&F University and collected in May, 2019, in order to investigate the changes in active ingredients and antioxidant capacity among different flowering stages (bud, initial, and full bloom). Changes in radical scavenging capability of DPPH (1,1-Diphenyl-2-picrylhydrazyl radical), ABTS (2,2′-azinobis(3-ethylbenzthiazoline-6-sulphonate)), and hydroxy were analyzed. Significant differences in active ingredient content of flowers were detected among different flowering stages. The total phenolic content increased continuously during the entire flowering stage. The contents of total flavonoid, total polysaccharide, and cyanidin-3-O-glucoside reached peaks at the initial blooming stage and then fell as the flowering process continued. The antioxidant activity in initial stage was the highest than in any other flowering stages. Therefore, we conclude that the initial blooming stage is the best harvesting stage of B. striata flowers. This study provides a robust basis for the harvest and utilization of B. striata flowers in food, medical, and cosmetic industries.