Pollination system of the Pilosocereus leucocephalus columnar cactus (tribe Cereeae) in eastern Mexico

It has been suggested that there is a geographic dichotomy in the pollination systems of chiropterophilous columnar cacti: in intra‐tropical areas they are pollinated almost exclusively by bats, whereas in extratropical areas they are pollinated by bats, birds and bees. However, currently the studies are clumped both taxonomically (mainly Pachycereeae species) and geographically (mainly in the Tehuacan Valley and the Sonoran Desert). This clumping limits the possibility of generalising the pattern to other regions or cactus tribes. Only four of the 36 chiropterophilous cacti in Pilosocereus have been studied. Despite the tropical distribution of two Pilosocereus species, bees account for 40–100% of their fruit set. We examined how specialised is the pollination system of P. leucocephalus in eastern Mexico. As we studied tropical populations, we expected a bat‐specialised pollination system. However, previous studies of Pilosocereus suggest that a generalised pollination system is also possible. We found that this cactus is mainly bat‐pollinated (bats account for 33–65% of fruit set); although to a lesser degree, diurnal visitors also caused some fruit set (7–15%). Diurnal visitors were more effective in populations containing honeybee hives. P. leucocephalus is partially self‐compatible (14–18% of fructification) but unable to set fruit without visitors. Despite the variation in pollination system, P. leucocephalus shows more affinity with other columnar cacti from tropical regions than with those from extratropical regions. Although we report here that a new species of tropical Pilosocereus is relatively bat‐specialised, this Cereeae genus is more flexible in its pollination system than the Pachycereeae genera.     

The nectar of the Strelitzia reginae flower

The nectar of Strelitzia reginae Ait. was analysed using enzymatic methods and found to contain glucose, fructose and sucrose. Sugar composition changed considerably over the nectar producing period: there was an increase in the amount of glucose (41%) and fructose (32%) between the early and middle stage of secretion and thereafter a decrease of 13 and 24%, respectively, towards the end of secretion. Although the amount of sucrose secreted was initially as much as the glucose and fructose combined, it subsequently decreased, first by 14% and then by 70% at the end of the secretory period so that, whereas in the initial stages of secretion sucrose was quantitatively the dominant sugar, glucose and fructose made up the major part of the nectar as secretion reached its conclusion.
The amounts of potassium and sodium remained at the same low level (around 150 and 30 μg g^-1 [w/v) respectively) throughout secretion, while calcium (initially 18 μg g^-1) and magnesium (initially 8.0 μg g^-1) increased by 47 and 56% respectively, between the early and late stages of secretion. No free amino acids, inorganic phosphate or iron could be detected. Enzymatic analysis revealed only a trace amount of starch. Transmission electron micrographs from both immature and mature plants, however, showed starch grains among other cytoplasmic remnants in the nectary lumen. Mitochondria, vesicles, lipid droplets and ribosomes could also be identified among the luminal cytoplasmic remnants.     

Pollination Ecology and Effect of Nectar Removal in Macleania bullata (Ericaceae)1

The floral syndrome of Macleania bullataYeo (Ericaceae) reflects its adaptation to hummingbird pollination. Its flowers, however, are subject to high levels of nectar robbing. I examined the floral visitor assemblage of M. bullata in a tropical montane wet forest in southwestern Colombia, focusing on the behavior of the visitors. I also tested for the presence of nocturnal pollination and the effects of nectar removal on new nectar production. The principal floral visitors were the nectar robbing hummingbirds Ocreatus underwoodii (19.1% of visits) and Chlorostilbon mellisugus (18.9%). Only two species of long–billed hummingbirds visited the flowers of M. bullata as “legitimate” pollinators: Coeligena torquata (14.7% of visits) and Doryfera ludoviciae (14.3%). The remaining visits constituted nectar robbing by bees, butterflies, and other species of hummingbirds. Nocturnal pollination took place, although fruit set levels were 2.4 times higher when only diurnal pollination was allowed as opposed to exclusively nocturnal pollination. Nectar robbers removed floral nectar without pollinating the flower. Treatments of experimental nectar removal were carried out to examine if flowers synthesize more nectar after nectar removal. Nectar removal increased the total volume of nectar produced by each flower without affecting sugar concentration. Thus, nectar robbing can impose a high cost to the plants by forcing them to replace lost nectar.     

Variability of nectar production and composition in Linaria vulgaris (L.) Mill. (Scrophulariaceae)

 We studied nectar characteristics during the long flowering period (late June to end of November) in two populations of Linaria vulgaris (L.) Mill. spontaneously growing in the Botanical Gardens of Siena University (Tuscany, central Italy). The two populations were close to each other but they differed in blooming period. Plants of population 1 sprouted in May and flowered from the end of June to the end of September. Population 2 sprouted at the end of August and flowered from September to the end of November. Differences in nectar production and composition were found between and within populations. Flowers of population 1 produced a very small amount of nectar (not collectable) that remained on the nectary surface. The quantity of nectar increased in late September, when each flower produced 2–3 μl of nectar that flowed into the spur. Total sugar concentration was 175.8 mg/ml in young flowers. Flowers of population 2 produced 5–8 μl of nectar with a total sugar concentration of 200.9 mg/ml in the young stage. In bagged senescent flowers nectar volume decreased in both populations and nectar sugar concentration decreased down to 11.6 mg/ml in population 2 and increased up to 289.6 mg/ml in population 1. For both populations, the decrease in nectar volume in bagged flowers may have been due to water loss by evaporation. In population 2, the decrease in sugar concentration may have been due to nectar reabsorption that was never observed in population 1. Nectar variability is discussed in relation to insect visits and seed set. Received August 14, 2002; accepted December 17, 2002 Published online: June 2, 2003     

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.     

Bumblebee Pollination of Melampyrum ciliare (Scrophulariaceae)

Abstract Melampyrum ciliare has tubular flowers predominantly visited by Bombus diversus. Floral structure (including the positions of anthers and stigma) and structure of the distal part of the corolla indicate cross-pollination by bumblebees. In M. ciliare, young flowers with white spots on the labium, which disappeared with flower aging, produced larger amounts of nectar than older ones without spots. Bumblebees visited flowers with white spots significantly more frequently than would be expected if they chose flowers randomly. These findings and the high seed production of open-pollinated flowers suggest effective pollination of M. ciliare by bumblebees.     

Floral nectar production: what cost to a plant?

Floral nectar production is central to plant pollination, and hence to human wellbeing. As floral nectar is essentially a solution in water of various sugars, it is likely a valuable plant resource, especially in terms of energy, with plants experiencing costs/trade-offs associated with its production or absorption and adopting mechanisms to regulate nectar in flowers. Possible costs of nectar production may also influence the evolution of nectar volume, concentration and composition, of pollination syndromes involving floral nectar, and the production of some crops. There has been frequent agreement that costs of floral nectar production are significant, but relevant evidence is scant and difficult to interpret. Convincing direct evidence comes from experimental studies that relate either enhanced nectar sugar production (through repeated nectar removal) to reduced ability to produce seeds, or increased sugar availability (through absorption of additional artificial nectar) to increased seed production. Proportions of available photosynthate allocated by plants to nectar production may also indicate nectar cost. However, such studies are rare, some do not include treatments of all (or almost all) flowers per plant, and all lack quantitative cost–benefit comparisons for nectar production. Additional circumstantial evidence of nectar cost is difficult to interpret and largely equivocal. Future research should repeat direct experimental approaches that relate reduced or enhanced nectar sugar availability for a plant with consequent ability to produce seeds. To avoid confounding effects of inter-flower resource transfer, each plant should experience a single treatment, with treatment of all or almost all flowers per plant. Resource allocation by plants, pathways used for resource transfer, and the locations of resource sources and sinks should also be investigated. Future research should also consider extension of nectar cost into other areas of biology. For example, evolutionary models of nectar production are rare but should be possible if plant fitness gains and costs associated with nectar production are expressed in the same currency, such as energy. It should then be possible to understand observed nectar production for different plant species and pollination syndromes involving floral nectar. In addition, potential economic benefits should be possible to assess if relationships between nectar production and crop value are evaluated.     

Does competition with wind‐pollinated species alter Echium plantagineum's attractiveness to a common pollinator Bombus terrestris?

1. In insect‐pollinated plants, pollinator attraction is influenced by flowers (e.g. number, size) and their associated rewards (e.g. pollen, nectar). These traits can depend on plant interactions. Indeed, below‐ground competition between plants can lead to a decrease in flower or reward production in insect‐pollinated species. 2. Wind‐pollinated plants, in particular, which are almost never studied in plant–pollinator networks, can alter insect‐pollinated plants' attractiveness through competition for nutrients. The response of pollinators to such changes has never been investigated. 3. A pot experiment was carried out in which an insect‐pollinated species, Echium plantagineum, was grown in binary mixture with three wind‐pollinated species selected to exert a panel of competitive interactions. Below‐ground competition was controlled using dividers limiting interspecific root competition. Floral traits of E. plantagineum (i.e. flower production, floral display size, flower size and nectar production) were measured. For each species mixture, the visits (i.e. first visit, number of visits, 10‐min sequences) of Bombus terrestris individuals released in a flight cage containing two pots were followed, one with and one without below‐ground competition. 4. Below‐ground competition significantly affected nectar's sucrose concentration but did not influence flower and nectar production. Likewise, pollinator visits were not influenced by below‐ground competition. Competitor identity significantly influenced flower and reward production of E. plantagineum, with a decrease in the presence of the most competitive wind‐pollinated species. A tendency for faster flower visitation events was also detected in the presence of the least competitive competitor. This study raises new questions regarding the influence of wind‐pollinated plants on plant–pollinator interactions.     

传粉昆虫对我国中南地区油茶结实和结籽的作用

油茶(Camellia oleifera)是一种重要的木本油料植物。2007-2008年在我国中南地区湖南省对油茶开展传粉生物学研究结果表明:油茶花期从10月到翌年1月份,单株开花时间一般持续20d左右,单花花期为4-5d。单花花蜜量平均为(145.40±24.89)μL,含糖量为(23.13±1.03)%,并含有17种氨基酸,约占花蜜总量的1.73%。膜翅目蜜蜂总科(Hymenoptera:Apoidea)油茶地蜂(Andrena camellia)和大分舌蜂(Colletes gigas)是油茶的有效传粉昆虫,大分舌蜂的传粉效率较油茶地蜂高;蝇类和蝶类为随机访问者。油茶不存在无融合生殖和自动自花授粉现象,其结实和结籽依赖传粉者。此外,油茶具有一定程度的自交不亲和,自交和异交的繁殖成功率存在显著性差异。人工异花授粉结果显示油茶在资源限制程度较低时,即大年时,花粉限制主要表现在结籽水平,而不是结实水平。根据油茶的传粉综合特征以及油茶在山茶属中的系统位置,目前很难解释油茶类似鸟类传粉综合特征的花蜜特征是如何演化的。     

传粉生物学中几种花蜜采集和糖浓度测定方法的比较

花蜜的研究是花生物学中的一个重要内容,探寻实用的方法将方便野外操作。我们分别还用毛细管、注射器、滤纸条和离心法采集了5种花的花蜜,以比较各种方法的优劣,并用3种旋光测糖仪测量了慈姑Sagittaria trifolia L,的雌雄花的花蜜糖含量。目的是为寻找一种适合小型花的花蜜采集测量方法。结果表明,几种方法的适用性受花的大小、形状、蜜的分泌量及蜜腺位置的影响非常大,不同的花要采用不同的方法。对于一般的野外工作建议用毛细管采集后使用便携式旋光测糖仪测其糖含量。特别小的花和蜜量微小的花可以采用离心法收集。