Small but attractive: female-biased nectar production and floral visitors in a dimorphic shrub.

In sexually dimorphic species, hermaphrodite flowers in gynodioecious species or male flowers in dioecious species are often larger and produce more nectar than their conspecific female flowers. As a consequence, hermaphrodite or male flowers frequently receive more pollinator visits. Sex ratio, flower size, floral display, nectar production and floral visits were evaluated in two natural populations of Fuchsia thymifolia, a morphologically gynodioecious but functionally subdioecious insect-pollinated shrub. Sex ratio did not differ from the expected 1:1 in the two studied populations. As expected, hermaphrodite flowers were larger than female flowers, but in contrast to the general pattern, hermaphrodite flowers did not produce nectar or produced much less than female flowers. Flower visitors were flies (68%) and bumblebees (24%), both of which showed a preference for female flowers. No sex difference was detected in either flower longevity or floral display across the flowering season. Higher nectar production by females may attract more pollinators, and may be a strategy to enhance female reproductive success in this species. Finally, floral dimorphism and insect preferences did not seem to hamper the maintenance of sub-dioecy or prevent the evolution of dioecy in F. thymifolia.

Differences in Copper Absorption and Accumulation between Copper-Exclusion and Copper-Enrichment Plants: A Comparison of Structure and Physiological Responses.

Differences in copper (Cu) absorption and transport, physiological responses and structural characteristics between two types of Cu-resistant plants, Oenothera glazioviana (Cu-exclusion type) and Elsholtzia haichowensis (Cu-enrichment type), were investigated in the present study. The results indicated the following: (1) After 50 µM Cu treatment, the Cu ratio in the xylem vessels of E. haichowensis increased by 60%. A Cu adsorption experiment indicated that O. glazioviana exhibited greater resistance to Cu, and Cu absorption and the shoot/root ratio of Cu were significantly lower in O. glazioviana than in E. haichowensis. (2) An analysis of the endogenous abscisic acid (ABA) variance and exogenous ABA treatment demonstrated that the ABA levels of both plants did not differ; exogenous ABA treatment clearly reduced Cu accumulation in both plants. (3) The leaf stomatal density of O. glazioviana was significantly less than that of E. haichowensis. Guard cells in E. haichowensis plants were covered with a thick cuticle layer, the epidermal hair was more numerous and longer, and the number of xylem conduits in the root was small. (4) The transpiration rate and the stomatal conductance of O. glazioviana were both significantly lower than those of E. haichowensis, regardless of whether the plants were treated with Cu. Taken together, these results indicate that the differences in the structural characteristics between these two plant species, particularly in the characteristics related to plant transpiration, are important factors that govern whether plants acquire or exclude Cu.

Sex-specific reproductive components and pollination ecology in the subdioecious shrub Fuchsia microphylla.

In subdioecious populations, functional female, male and hermaphrodite individuals coexist. Subdioecy may be a transitional state towards dioecy or a breakdown of dioecy, although lability in sex expression may maintain subdioecy as a stable condition. To better understand the ecological aspects involved in sex ratio dynamics and breeding system evolution, we studied the pollination and female fitness components of female and hermaphrodite individuals of the subdioecious shrub Fuchsia microphylla. In two natural populations at the Trans-Mexican Volcanic Belt we estimated female frequency and several reproductive components of female and hermaphrodite plants under natural pollination and experimental pollination treatments. Average female frequency was 42%, and on average, 42.5% of hermaphrodites produced fruits. Female plants showed a 17-fold female fertility advantage over hermaphrodites through increased fruit production, as the number of seeds and germination rates did not differ between morphs. Hermaphrodite flowers were larger, with similar nectar production and concentration to female flowers, and pollinators did not show consistent morph preferences. Some hermaphrodites produced fruits autonomously, and female flowers excluded from pollinators produced fruits putatively by apomixis. Fruit production in hermaphrodites, but not in females, was related to height, suggesting increased investment of hermaphrodites in the female function at higher resource status. For sex ratios to be at equilibrium, the female fertility advantage should be reduced about eightfold. However, it may be that hermaphrodites are maintained by producing fruits at no cost to the male function at higher resource status, as the gender plasticity hypothesis proposes.

A fossil Fuchsia (Onagraceae) flower and an anther mass with in situ pollen from the early Miocene of New Zealand.

PREMISE OF THE STUDY: Fuchsia (Onagraceae) anthers, pollen, and an ornithophilous Fuchsia-like flower from an earliest Miocene lacustrine diatomite deposit at Foulden Maar, southern New Zealand confirm a long record for Fuchsia in New Zealand and probably an equally long history for its distinctive honeyeater pollination syndrome. The anthers contain in situ pollen of the fossil palynomorph previously assigned to Diporites aspis Pocknall et Mildenh. (Onagraceae: Fuchsia L.). • METHODS: We undertook comparative studies of the flower and anther morphology of the newly discovered macrofossils and compared the in situ pollen grains from the anthers with dispersed pollen grains from extant species. • KEY RESULTS: The anther mass is referred to a new, extinct species, Fuchsia antiqua D.E.Lee, Conran, Bannister, U.Kaulfuss & Mildenh. (Onagraceae), and is associated with a fossilized Fuchsia-like flower from the same small mining pit. Because Diporites van der Hammen is typified by a fungal sporomorph, the replacement name for D. aspis is Koninidites aspis (Pocknall & Mildenh.) Mildenh. gen. & comb. nov. Phylogenetic placement of the fossils agrees with a proximal position to either sect. Skinnera or sect. Procumbentes. These are the oldest macrofossils of Fuchsia globally. • CONCLUSIONS: The floral structures are remarkably similar to those of modern New Zealand Fuchsia. They suggest that the distinctive honeyeater bird-pollination syndrome/association seen in modern New Zealand was already established by the late Oligocene-earliest Miocene. The implications for the biogeography and paleoecology of Fuchsia in Australasia are discussed.

Attracting pollinators and avoiding herbivores: insects influence plant traits within and across years.

Perennial plants interact with herbivores and pollinators across multiple growing seasons, and thus may respond to herbivores and pollinators both within and across years. Joint effects of herbivores and pollinators influence plant traits, but while some of the potential interactions among herbivory, pollination, plant size, and plant reproductive traits have been well studied, others are poorly understood. This is particularly true for perennial plants where effects of herbivores and pollinators may manifest across years. Here, we describe two experiments addressing the reciprocal interactions of plant traits with herbivore damage and pollination across 2 years using the perennial plant Chamerion angustifolium. We measured (1) plant responses to manipulation of damage and pollination in the year of treatment and the subsequent season, (2) damage and pollination responses to manipulation of plant size and flowering traits in the year of treatment, and (3) plant-mediated indirect interactions between herbivores and pollinators. We found that plant traits had little effect on damage and pollination, but damage and pollination affected plant traits in both the treatment year and the subsequent year. We found evidence of indirect effects between leaf herbivores and pollinators in both directions; indirect effects of pollinators on leaf herbivores have not been previously demonstrated. Our results indicate that pollen receipt results in shorter plants with fewer stems but does not change flower number, while leaf herbivory results in taller plants with fewer flowers. Together, herbivory and pollination may contribute to intermediate plant height and plants with fewer stems and flowers in our system.

Mechanics of plant fruit hooks.

Hook-like surface structures, observed in some plant species, play an important role in the process of plant growth and seed dispersal. In this study, we developed an elastic model and further used it to investigate the mechanical behaviour of fruit hooks in four plant species, previously measured in an experimental study. Based on Euler-Bernoulli beam theory, the force-displacement relationship is derived, and its Young's modulus is obtained. The result agrees well with the experimental data. The model aids in understanding the mechanics of hooks, and could be used in the development of new bioinspired Velcro-like materials.

Precise spatio-temporal regulation of the anthocyanin biosynthetic pathway leads to petal spot formation in Clarkia gracilis (Onagraceae).

Petal spots are widespread in angiosperms and are often implicated in pollinator attraction. Clarkia gracilis petals each have a single red-purple spot that contrasts against a pink background. The position and presence of spots in C. gracilis are determined by the epistatic interaction of alleles at two as yet unidentified loci. We used HPLC to identify the different pigments produced in the petals, and qualitative and quantitative RT-PCR to assay for spatio-temporal patterns of expression of different anthocyanin pathway genes. We found that spots contain different pigments from the remainder of the petal, being composed of cyanidin/peonidin-based, instead of malvidin-based anthocyanins. Expression assays of anthocyanin pathway genes showed that the dihydroflavonol-4-reductase 2 (Dfr2) gene has a spot-specific expression pattern and acts as a switch for spot production. Co-segregation analyses implicated the gene products of the P and I loci as trans-regulators of this switch. Spot pigments appear earlier in development as a result of early expression of Dfr2 and the flavonoid 3' hydroxylase 1 (F3'h1) gene. Pigments in the background appear later, as a result of later expression of Dfr1 and the flavonoid 3'-5' hydroxylase 1 (F3'5'h1) genes. The evolution of this spot production mechanism appears to have been facilitated by duplication of the Dfr gene and to have required substantial reworking of the anthocyanin pathway regulatory network.

Genome duplication and the evolution of physiological responses to water stress.

Whole-genome multiplication, or polyploidy, is common in angiosperms and many species consist of multiple cytotypes that have different physiological tolerances. However, the relative importance of genome duplication vs post-duplication evolutionary change in causing differentiation between cytotypes is not known. We examined the water relations of Chamerion angustifolium, a herbaceous perennial in which diploid and tetraploid cytotypes occupy different niches. To differentiate between the effects of genome duplication and evolutionary changes that followed polyploidization, we compared extant diploids and tetraploids with experimentally synthesized neotetraploids. Tetraploids had 32% higher xylem hydraulic conductivity (K(H)) than neotetraploids and 87% higher K(H) than diploids, but vulnerability to water stress induced cavitation and gas exchange sensitivity to water potential did not differ among cytotypes. Nevertheless, tetraploids took 22% and 30% longer to wilt than neotetraploids and diploids. A simple hydraulic model suggested that tetraploids deplete soil moisture to a greater degree than neotetraploids and diploids before reaching leaf water potentials that cause stomatal closure. We conclude that the different physiological tolerances and distribution of diploid and tetraploid C. angustifolium are unlikely to be caused solely by genome duplication. The enhanced ability of tetraploids to survive water stress likely evolved after polyploidization.