Outstanding questions in flower metabolism

The great diversity of flowers, their color, odor, taste, and shape, is mostly a result of the metabolic processes that occur in this reproductive organ when the flower and its tissues develop, grow, and finally die. Some of these metabolites serve to advertise flowers to animal pollinators, other confer protection towards abiotic stresses, and a large proportion of the molecules of the central metabolic pathways have bioenergetic and signaling functions that support growth and the transition to fruits and seeds. Although recent studies have advanced our general understanding of flower metabolism, several questions still await an answer. Here, we have compiled a list of open questions on flower metabolism encompassing molecular aspects, as well as topics of relevance for agriculture and the ecosystem. These questions include the study of flower metabolism through development, the biochemistry of nectar and its relevance to promoting plant‐pollinator interaction, recycling of metabolic resources after flowers whiter and die, as well as the manipulation of flower metabolism by pathogens. We hope with this review to stimulate discussion on the topic of flower metabolism and set a reference point to return to in the future when assessing progress in the field.     

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.     

Nectar concentration and composition of 26 species from the temperate forest of South America

BACKGROUND AND AIMS: Floral nectar concentration and chemical composition of 26 plant species native to the temperate forest of southern South America are reported and the relationships with the flower type are evaluated. METHODS: Nectar concentration was measured with a hand refractometer and sugar composition was analysed by gas-liquid chromatography. Plant species were classified into flower type categories based not only on floral features but also on data from the literature and field observations on their pollinators. KEY RESULTS: Most data on nectar are new reports at the generic and/or specific level. Plant species in which more than one population was studied showed significant among-population variation in nectar sugar concentration and composition. Results showed a weak relationship between nectar traits and flower type. Many species had nectar containing 50 % or more sucrose (17 of 26 species), independent of the main pollinator. CONCLUSIONS: Considering that (a) nectar characteristics did not show a clear association with different flower types or with plant taxonomic membership, and (b) different populations of the same species showed large variability in sugar composition, the results suggest that other factors (e.g. historical and environmental) could be involved in determining the sugar composition of the highly endemic plant species from this region.     

Nectar Dynamics and Sugar Composition in Flowers of Silene and Saponaria Species (Caryophyllaceae)

Abstract: Nectar production in Saponaria officincilis and in five species of Silene (S. ciba, S. dioica, S. noctiflora, S. nutans, S. vulgaris ) was examined during two consecutive years (May to July 1993, and May to June 1994) in the Botanical Garden of the University of Giessen. Nectar volume and sugar concentration were studied in relation to time of day, flower sex, flower age, and flowering stage. Nectar amount in all species studied (except S. dioica ) increased in the afternoon or in the evening until midnight (or until the early morning in S. nutans ). After midnight and until midday, nectar volume in non-visited flowers (except S. dioica ) decreased. Nectar volume in non-visited S. dioica flowers increased constantly with flower age, indicating a stable nectar secretion rate, possibly favouring both day- and night-active flower visitors. Even at the time of highest nectar secretion, all species studied presented several nectarless flowers. Sucrose dominance in the nectar of the nocturnal species S. nutans and Saponaria officinalis fits well with the general syndrome of flowers pollinated by hawkmoths. The syndrome also applies to the nocturnal but regularly selfing, S. noctiflora . The more generalis-tic species S. dioica and S. vulgaris , which are regularly visited by bumblebees as well as nocturnal moths, secreted hexose-domi-nant nectar. Unexpectedly, Silene alba , the only nocturnal species that strictly excluded day-active flower visitors by closing flowers during the day, also secreted hexose-dominant nectar. In some cases, nectar volumes and nectar concentration differed significantly between hermaphroditic, male, and female flowers. Female flowers of S. alba, S. dioica , and S. nutans contained significantly less concentrated nectar than male or hermaphroditic ( S. nutans ) ones. In S. noctifiora and S. vulgaris the difference was not statistically significant but nectar concentration did show the same tendency.     

Rewardlessness in orchids: how frequent and how rewardless?

About one‐third of orchid species are thought to offer no floral reward and therefore attract pollinators through deception. Statements of this idea are common in the botanical literature, but the empirical basis of the estimate is rarely mentioned. We traced citation pathways for the one‐third estimate in a sample of the literature and found that the paths lead to empirical foundations that are surprisingly narrow. Moreover, recent measurements have detected minute quantities of sugar available to insect visitors in some orchids thought to be rewardless, raising the possibility of a pollination strategy that is largely deceitful but different to absolute rewardlessness. The orchids are a well‐studied group and there is no doubt that rewardlessness is common in the family. However, greater empirical effort is needed to verify rewardlessness in orchids and to explore geographic and environmental variation in the proportion of rewardless species.