Floral Advertisement Research Articles

Floral scent in a whole‐plant context: moving beyond pollinator attraction

The natural world is awash with odours – blends of volatile organic compounds (VOCs) – whose chemical structures and composition convey information to organisms endowed with the sensory capacity to perceive them. Pheromones and other chemical signals provide specific information that facilitates kin recognition, mate acquisition, social interactions and defence across the tree of life (Meinwald & Eisner 2008). Even more general volatile cues, such as CO2, convey context-dependent information about habitat and host or food quality, illness and microbial decay (Guerenstein & Hildebrand 2007). The terrestrial fossil record teems with evidence of chemical communication and warfare, in the form of animal olfactory appendages (Strausfeld & Hildebrand 1999; Labandeira 2002) and plant essential oil glands (Fahn 2002; Krings, Taylor & Kellogg 2002). This evidence, combined with our knowledge of cycads, water lilies and other ‘living fossils’ (Thien, Azuma & Kawano 2000; Terry et al. 2007), attests to the antiquity of volatile communication between plants, their mutualists and antagonists.

FLORAL BIOLOGY OF NORTH AMERICAN OENOTHERA SECT. LAVAUXIA (ONAGRACEAE): ADVERTISEMENTS, REWARDS, AND EXTREME VARIATION IN FLORAL DEPTH1,2

We studied the floral biology of five North American members of Oenothera L. sect. Lavauxia (Spach) Endl. (Onagraceae L.) in field and common greenhouse settings. Oenothera sect. Lavauxia floral morphology ranges from small, cleistogamous flowers (O. flava subsp. flava (A. Nels.) Garrett in Garrett) to some of the longest-tubed flowers in North America (O. flava subsp. taraxacoides (Wooton & Standl.) W. L. Wagner). Our goal was to compare qualitative and quantitative aspects of floral advertisement and reward among taxa in section Lavauxia. All taxa are night-blooming and self-compatible, have yellow petals with ultraviolet reflectance, and produce floral scents dominated by nitrogenous compounds and monoterpenes. Methyl nicotinate is present in the fragrances of all taxa of section Lavauxia regardless of flower size or putative mating system. Because this floral volatile is largely absent from other Oenothera species, we hypothesize that it is a synapomorphy for section Lavauxia. The rare O. acutissima W. L. Wagner, which is endemic to the Uintah Mountains, is polymorphic for odors dominated by linalool- or ocimene-derived compounds. Field observations in its type locality in northeastern Utah, U.S.A., revealed frequent floral visitation by crepuscular hawkmoths during the first 1.5 hours after anthesis, a pattern common to O. flava subsp. taraxacoides and other large-flowered Oenothera throughout western North America. Quantitative aspects of floral advertisement (flower size, scent emission) and reward (nectar volume) are dramatically reduced in putatively autogamous taxa (O. flava subsp. flava, O. triloba Nutt.), whereas qualitative aspects (flower color, scent, and nectar chemistry) remain comparable. All taxa could be distinguished through ordination of characters related to flower size, herkogamy, and scent chemistry. Extreme nectar tube length variation across the range of O. flava renders this an excellent model system for measuring the costs and mechanisms of shifts between outcrossing and autogamy.

The Effect Of Pollination On Floral Fragrance in Thistles

We investigated postpollination changes in fragrance composition and emission rates, as well as pollinator discrimination in hand-pollinated flower heads of two thistle species: Canada thistle (Cirsium arvense) and sandhill thistle (C. repandum). Following pollination, neither species emitted any novel compounds that could function as repellents. Scent emission rates declined in pollinated plants of both species by approximately 89% within 48 hr. This decline was evident in all 13 scent components of C. arvense. Apis mellifera, the dominant pollinator in the study population of C. arvense, was nearly three times more likely to visit an unpollinated rather than a pollinated flower head. A more complex pattern was observed for C. repandum, whose scent comprised 42 compounds. Quantities of aromatic and sesquiterpenoid volatiles declined after pollination, whereas two classes of scent compounds, fatty acid derivatives and monoterperpenoids, continued to be emitted. In C. repandum, discrimination against pollinated flower heads by Papilio palamedes (its primary pollinator) was not as marked. Unpollinated control plants of both species maintained moderate levels of scent production throughout this experiment, demonstrating that senescence and floral advertisement may be delayed until pollination has occurred. We expect postpollination changes in floral scent contribute to communication between plants with generalized pollinator spectra and their floral visitors. This study provides the first field study of such a phenomenon outside of orchids.

Flowers as sensory billboards: progress towards an integrated understanding of floral advertisement.

This paper explores the landscape of integrated sensory signals that are produced by flowers, and the contextual information that modulates pollinator responses to such signals. Chiloglottis orchids are pollinated by mimicking the pheromone and posture of female thynnine wasps, but floral height provides the context within which male wasps respond to these signals. The odor and appearance of carrion attract blowflies to Helicodiceros inflorescences, but flies are more likely to enter the floral chamber when heat is present as a contextual cue. Finally, fragrance, UV-visual cues and echo fingerprints are redundant signals that Glossophaga bats can use to find flowers, depending on prior experience and the photic environment.

EXAPTATIONS LINK EVOLUTION OF PLANT–HERBIVORE AND PLANT–POLLINATOR INTERACTIONS: A PHYLOGENETIC INQUIRY

To evaluate possible evolutionary links between plant–herbivore and plant–pollinator relationships, defense and reward characteristics and pollination ecology were mapped onto a morphologically estimated phylogeny of 42 species of Dalechampia. This procedure generated detailed hypotheses about the evolution of anti-herbivore defense and pollination systems. These hypotheses were tested using the results of chemical analyses and bioassays of foliar and floral secretions. Multiple lines of defense appear to have evolved in sequence in Dalechampia. The first defense system to originate was deployment of triterpene resins to defend the staminate flowers. This feature was a preaptation (preadaptation) that allowed the evolution of a resin-based, pollinator-reward system. Thus pollination by resin-collecting bees originated as a “transfer exaptation” (a new function replaced the old). This hypothesis is supported by anti-herbivore activities of floral resins and by chemical similarity of floral defense and reward resins. After the resin defense of flowers was lost by conversion into a reward system, there followed (in evolutionary time) a sequence of defensive innovations. These included nocturnal closure of large, involucral bracts to protect both staminate and pistillate flowers. Phylogenetic analysis showed that the large bracts themselves probably originated as a floral advertisement system, and the bracts assumed a defensive function through “addition exaptation” (a new function was added to the old). Additional lines of defense to evolve were deployment of resin to defend the developing ovaries and seeds (an addition exaptation), deployment of sharp, detaching trichomes on enveloping sepals to defend developing seeds (apparent adaptations), closure of involucral bracts around the developing fruits and seeds (an addition exaptation), and deployment of resin to defend the leaves and growing shoot tips (also an addition exaptation). Support for this scenario also derives from the chemical similarities of sepal, foliar, and floral resins, and the anti-herbivore properties of foliar resins. It appears that at least one pollinator-reward system originated by modification of a defense system, and several defense systems originated by modification of pollinator reward and advertisement systems. Thus exaptations have been important in the origin of new defense and pollination systems, and each system has significantly influenced the evolution of the other on several occasions.

SIZE AND SHAPE IN FLORAL ADVERTISEMENT: MEASUREMENT, CONCEPTS AND IMPLICATIONS

SIZE AND SHAPE IN FLORAL ADVERTISEMENT: MEASUREMENT, CONCEPTS AND IMPLICATIONS

Bateman’s principle and plant reproduction: The role of pollen limitation in fruit and seed set

Bateman’s principle states that male fitness is usually limited by the number of matings achieved, while female fitness is usually limited by the resources available for reproduction. When applied to flowering plants this principle leads to the expectation that pollen limitation of fruit and seed set will be uncommon. However, if male searching for mates (including pollen dissemination via external agents) is not sufficiently successful, then the reproductive success of both sexes (or both sex functions in hermaphroditic plants) will be limited by number of matings rather than by resources, and Bateman’s principle cannot be expected to apply. Limitation of female success due to inadequate pollen receipt appears to be a common phenomenon in plants. Using published data on 258 species in which fecundity was reported for natural pollination and hand pollination with outcross pollen, I found significant pollen limitation at some times or in some sites in 159 of the 258 species (62%). When experiments were performed multiple times within a growing season, or in multiple sites or years, the statistical significance of pollen limitation commonly varied among times, sites or years, indicating that the pollination environment is not constant. There is some indication that, across species, supplemental pollen leads to increased fruit set more often than increased seed set within fruits, pointing to the importance of gamete packaging strategies in plant reproduction. Species that are highly self-incompatible obtain a greater benefit relative to natural pollination from artificial application of excess outcross pollen than do self-compatible species. This suggests that inadequate pollen receipt is a primary cause of low fecundity rates in perennial plants, which are often self-incompatible. Because flowering plants often allocate considerable resources to pollinator attraction, both export and receipt of pollen could be limited primarily by resource investment in floral advertisement and rewards. But whatever investment is made is attraction, pollinator behavioral stochasticity usually produces wide variation among flowers in reproductive success through both male and female functions. In such circumstances the optimal deployment of resources among megaspores, microspores, and pollinator attraction may often require more flowers or more ovules per flower than will usually be fertilized, in order to benefit from chance fluctuations that bring in large number of pollen grains. Maximizing seed set for the entire plant in a stochastic pollination environment might thus entail a packaging strategy for flower number or ovule number per flower that makes pollen limitation of fruit or seed set likely. Pollen availability may limit female success in individual flowers, entire plants (in a season or over a lifetime), or populations. The appropriate level must be distinguished depending on the nature of the question being addressed.

Genetic systems of six species ofPlantago (Plantaginaceae)

Investigation of the genetic system of six species ofPlantago has revealed striking differences in their breeding and meiotic systems.Plantago patagonica is an inbreeder on account of preanthesis cleistogamy, whereasP. lanceolata is an obligate outbreeder, as it is self-incompatible.Plantago drummondii, P. lagopus, P. ovata, andP. major show mixed mating but in varying proportions. In terms of their energy budgets, outbreeding species invest more in floral advertisement and male function, while inbreeders invest more in female function. The contribution of the meiotic system to genetic variability, as revealed through recombination index, is more important in the inbreeding species.