Floral traits mediate the vulnerability of aloes to pollen theft and inefficient pollination by bees

Background and Aims

Pollen-collecting bees are among the most important pollinators globally, but are also the most common pollen thieves and can significantly reduce plant reproduction. The pollination efficiency of pollen collectors depends on the frequency of their visits to female(-phase) flowers, contact with stigmas and deposition of pollen of sufficient quantity and quality to fertilize ovules. Here we investigate the relative importance of these components, and the hypothesis that floral and inflorescence characteristics mediate the pollination role of pollen collection by bees.

Methods

For ten Aloe species that differ extensively in floral and inflorescence traits, we experimentally excluded potential bird pollinators to quantify the contributions of insect visitors to pollen removal, pollen deposition and seed production. We measured corolla width and depth to determine nectar accessibility, and the phenology of anther dehiscence and stigma receptivity to quantify herkogamy and dichogamy. Further, we compiled all published bird-exclusion studies of aloes, and compared insect pollination success with floral morphology.

Key Results

Species varied from exclusively insect pollinated, to exclusively bird pollinated but subject to extensive pollen theft by insects. Nectar inaccessibility and strong dichogamy inhibited pollination by pollen-collecting bees by discouraging visits to female-phase (i.e. pollenless) flowers. For species with large inflorescences of pollen-rich flowers, pollen collectors successfully deposited pollen, but of such low quality (probably self-pollen) that they made almost no contribution to seed set. Indeed, considering all published bird-exclusion studies (17 species in total), insect pollination efficiency varied significantly with floral shape.

Conclusions

Species-specific floral and inflorescence characteristics, especially nectar accessibility and dichogamy, control the efficiency of pollen-collecting bees as pollinators of aloes.     

The gene expression landscape of thermogenic skunk cabbage suggests critical roles for mitochondrial and vacuolar metabolic pathways in the regulation of thermogenesis

Floral thermogenesis has been described in several plant species. Because of the lack of comprehensive gene expression profiles in thermogenic plants, the molecular mechanisms by which floral thermogenesis is regulated remain to be established. We examined the gene expression landscape of skunk cabbage (Symplocarpus renifolius) during thermogenic and post-thermogenic stages and identified expressed sequence tags from different developmental stages of the inflorescences using super serial analysis of gene expression (SuperSAGE). In-depth analysis suggested that cellular respiration and mitochondrial functions are significantly enhanced during the thermogenic stage. In contrast, genes involved in stress responses and protein degradation were significantly up-regulated during post-thermogenic stages. Quantitative comparisons indicated that the expression levels of genes involved in cellular respiration were higher in thermogenic spadices than in Arabidopsis inflorescences. Thermogenesis-associated genes seemed to be expressed abundantly in the peripheral tissues of the spadix. Our results suggest that cellular respiration and mitochondrial metabolism play key roles in heat production during floral thermogenesis. On the other hand, vacuolar cysteine protease and other degradative enzymes seem to accelerate senescence and terminate thermogenesis in the post-thermogenic stage.     

Biotic versus abiotic pollination in Oeceoclades maculata (Lindl.) Lindl. (Orchidaceae)

Geographical variation in the reproductive biology of widespread species often occurs at their distributional boundaries. We sought to determine whether such variation has occurred in an invasive orchid, Oeceoclades maculata, across its naturalized range. We compared its reproductive biology in a Brazilian population with that published for a population on the Caribbean island of Puerto Rico. In the state of São Paulo, O. maculata flowers between December and February, at the height of the rainy season. Similar fruit sets were observed in manual self (76%) and cross (70.4%) pollination treatments. The fruit set of plants protected from both pollinators and rainfall was 6.1%, whereas plants exposed only to rainfall had a fruit set of 41.4%, slightly less than the controls (48.3%). Like the Puerto Rico population, reproduction is primarily through rain‐assisted autogamy, but unlike observations made on the island, outcrossing can eventually occur. We observed two butterfly species (Heliconius ethilla narcaea and Heliconius erato phyllis) pollinating O. maculata. Secretory epidermal cells and trichomes of the spur lumen produced 0.7 µL of 25% (sucrose equivalents) nectar per flower each morning, which was stored in a dilated basal portion of the spur and reabsorbed by the afternoon. Thus, geographical variation in reproductive biology exists across the broad invasive range of O. maculata.     

Comparative floral structure and systematics in Ochnaceae s.l. (Ochnaceae,Quiinaceae and Medusagynaceae; Malpighiales)

Ochnaceae s.l. (Ochnaceae, Quiinaceae and Medusagynaceae), one of the well‐supported subclades of the large order Malpighiales retrieved so far in molecular phylogenetic studies, were comparatively studied with regard to floral structure using microtome section series and scanning electron microscopy (SEM). Floral morphology, anatomy and histology also strongly reflect this close relationship. Potential synapomorphies of the subclade include: flowers nectarless, sepals of different sizes within a flower, petals not retarded in development and forming the protective organs of advanced floral buds, petal aestivation contort, petals with three vascular traces, petals reflexed over the sepals and directed toward the pedicel, polystemony, anthers almost or completely basifixed, gynoecium often with more than five carpels, short gynophore present, styles separate for at least their uppermost part and radiating outwards, suction‐cup‐shaped stigmas, vasculature forming a dorsal band of bundles in the upper stylar region, gynoecium epidermis with large, radially elongate cells, ovules either weakly crassinucellar or incompletely tenuinucellar with an endothelium, abundance of tanniferous tissues and sclerenchyma in floral organs. The most strongly supported subclade of two of the three families in molecular analyses, Quiinaceae and Medusagynaceae, is also particularly well supported by floral structural features, including the presence of functionally and morphologically unisexual flowers, a massive thecal septum that persists after anther dehiscence, styles radiating outward from the ovary, two lateral ovules per carpel, positioned one above the other, conspicuous longitudinal ribs on the ovary wall at anthesis, and a ‘false endothelium’ on the nucellus at anthesis. Additionally, the group fits well in Malpighiales and further emphasizes the relationship of Malpighiales with Celastrales and Oxalidales, and thus the unity of the COM clade. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 170 , 299–392.     

Fly pollination in Cypripedium: a case study of sympatric C. sichuanense and C. micranthum

Most Cypripedium spp. are known to be pollinated by bees. However, myiophilous traits are found in some species, especially in sections Trigonopedia and Sinopedilum. Here we chose C. micranthum and C. sichuanense, two sympatric species endemic to Sichuan, China, to test whether these orchids are fly pollinated. Artificial pollination showed that both flowers are self‐compatible but need pollen vectors for successful reproduction. Field observation showed that C. micranthum was pollinated by fruit flies and C. sichuanense by dung flies, both novel pollinators of Cypripedium orchids. These sympatric Cypripedium spp. are also cross‐compatible, but hybrids were not found in nature. The pollination syndromes of C. sichuanense and C. micranthum fit into the complex sapromyiophily pattern. It appears that pollinator specificity is responsible for their reproductive isolation. The discovery of fly pollination in C. sichuanense and C. micranthum, which belong to the related sections Trigonopedia and Sinopedilum, suggests a shift from bee to fly pollination in the genus Cypripedium. Unlike most Cypripedium spp., the anthers of C. micranthum release discrete pollinia with narrow stalks instead of the usual amorphous pollen smears. This ‘proto‐pollinarium’ is described, probably for the first time. These pollinia are most likely an adaptation for pollination by microdiptera, so the fly can carry the contents of both chambers in the same anther. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 170 , 50–58.     

Effects of planting method on the recovery of arboreal ant activity on revegetated farmland

Ecological restoration aims to re‐establish both biodiversity and ecological function in damaged ecosystems. Ants are important drivers of ecological functions and are early colonizers of restored ecosystems. Rates at which ants perform functions are thought to be fuelled by access to plant sugars. In revegetated farmland in south‐eastern Australia, I tested if ant activity on trees, which reflects use of arboreal sugars, follows a predictable trajectory of recovery towards a remnant‐like state. Additionally, I examined whether planting method alters this trajectory by comparing tube stock (TS), which results in low Eucalyptus densities, with direct seeding (DS), which results in high Eucalyptus densities. Replicate sites (n = 5) of young (planted between 1998 and 2001) and old (planted between 1989 and 1994) TS and DS revegetation, pastures and remnants were compared. Activity on trunks was significantly positively correlated with ant tending of Hemiptera in young and old revegetation. In DS plantings, activity and estimated liquid loads on Eucalyptus trees were low and rapidly approached that in remnants, while TS sites remained similar to high values observed in pastures with trees. Patterns for Acacia were less clear, reflecting consistent densities for this species between TS and DS. At the whole‐of‐field scale, planting methods did not differ. Importantly, although trajectories differed, neither TS nor DS sites approached the low activity or estimated liquid loads observed in remnants. Rates of ant use of arboreal sugars and associated sugar‐fuelled processes may thus take considerably longer to recover than the period covered by this study. This finding suggests planting method may affect the trajectory and outcome of revegetation for plant health, as well as sugar‐fuelled ecosystem functions performed by ants.     

濒危药用植物云南黄连传粉生态学研究

云南黄连为中药材黄连的原植物之一,为国家二级濒危保护植物。通过对云南黄连的开花物候、花部特征、繁育系统及传粉方式进行观察研究,以探讨云南黄连濒危机制,为其种质资源保护及人工抚育奠定基础。结果表明:(1)云南黄连的花期从12月份起至第二年的3月份,花期长达4个月,花序开花可持续45～60d,同一花序上不同花的开花时间相隔1～3d,单花花期可持续40～45d。(2)云南黄连为多歧聚伞花序,苞片包被花芽,花两性,雄蕊和心皮多数。(3)云南黄连的花粉-胚珠比(P/O)为9 000左右,杂交指数(OCI)值为4或5,为自交亲和但需要传粉者完成传粉的兼性异交型繁育系统,并存在无融合生殖现象。     

Colored nectar as an honest signal in plant-animal interactions

Many flowering plants obtain the services of pollinators by using their floral traits as signals to advertise the rewards they offer to visitors—such as nectar, pollen and other food resources. Some plants use colorful pigments to draw pollinators’ attention to their nectar, instead of relying on the appeal of nectar taste. Although this rare floral trait of colored nectar was first recorded by the Greek poet Homer in the Odyssey, it has only recently received the attention of modern science. This mini-review focuses on recent findings about some of the species that use colored nectar; topics include its function as an honest signal for pollinators, as well as the pigments responsible for the nectar coloration. Such research of the ecology and physiology of colored nectar expands our understanding of the role and evolution of pollinator signaling in plants.