Variation of flower opening and closing times in F1 and F2 hybrids of daylily (Hemerocallis fulva; Hemerocallidaceae) and nightlily (H. citrina)

In flowering plants, pollination success is strongly dependent on the timing of when flowers start to bloom and when they start to close. To elucidate the genetic mechanism influencing the timing of flower opening and closing, we obtained F1 and F2 hybrids of Hemerocallis fulva (a diurnally blooming species, pollinated by swallowtail butterflies) and H. citrina (a nocturnally blooming species, pollinated by nocturnal hawkmoths) and observed their flowering behavior from blooming to closing with the use of digital cameras. For flower opening times, F1 hybrids were highly variable, and F2 hybrids showed a bimodal distribution of flower opening times with peaks in both the morning and evening. The ratio of morning flowering and evening flowering among F2 hybrids did not deviate from 1:1. For the start to close time, both F1 and F2 hybrids were similar in showing the major peak in the evening. The ratio of evening closing and morning closing among F2 hybrids did not deviate from 3:1. These results suggest that the time of flower opening and the start of closing are regulated by different major genes.     

No evidence for a role of pollinator discrimination in causing selection on flower size through female reproduction

The preference for certain floral phenotypes by flower visiting animals may fuel the evolution of floral traits because variation in flower visitation rates may lead to fitness variation within a population. Here, I examine the importance of flower size for pollinator visitation rate, seed set, and seed mass in two alpine populations of the insect-pollinated herb Ranunculus acris L. during two seasons. There was no pollen limitation of seed set or mass. Pollinators discriminated strongly against flowers experimentally reduced in size. Despite this, there were no signs of any significant impact of flower size on female reproductive success. The results show that although pollinators discriminate strongly among floral phenotypes, this may not always result in female fitness differences within a population because seed set or mass is not limited by pollen availability alone. Probably abiotic environmental constraints prevent plants with high pollinator visitation from capitalizing on the high pollen deposition.     

Collinsia sparsiflora in serpentine and nonserpentine habitats: using F2 hybrids to detect the potential role of selection in ecotypic differentiation

Here we document phenotypic differences between serpentine and nonserpentine ecotypes of Collinsia sparsiflora, as well as patterns of selection in these contrasting soil habitats. We transplanted the two parental ecotypes and experimental F2 hybrids into six field sites, and collected morphological, phenological and fitness data on emergent plants. To focus on edaphically mediated selection, rather than on pollinator-mediated selection, we used pollinator-exclusion cages. Transplanted parentals of the two ecotypes showed genetic differentiation for floral traits, but not for phenological traits or cotyledon size. For the F2 hybrids growing on serpentine soils, there was significant directional selection on cotyledon size, flower size and flower shape. However, the pattern of selection did not differ significantly between serpentine and nonserpentine sites. Overall, we did not see evidence for divergent selection on the two soil types. We conclude that differences in floral traits between the ecotypes do not reflect adaptation to physical conditions associated with soil type, and that there are unmeasured traits that must be contributing to ecotypic differentiation.     

An experimental test of hybrid resistance to insects and pathogens using Salix caprea, S. repens and their F1 hybrids

The aim of this study was to assess the responses of herbivores and pathogens to hybrid plants under controlled conditions. F1 hybrids and parental species, produced by hand-pollinating willows in the field, were potted and kept in an experimental field under controlled conditions. In 1997, plant growth and survival were measured along with densities of insects and the degree of pathogen infection on the willows. The survival rate was higher for S. repens than for the hybrids and lowest for S. caprea. Densities of the sawflies Pontania pedunculi and P. brigmanii and the leaf-galling midge Iteomyia capreae were higher on hybrids and on S. caprea than on S. repens. The densities of Crepidodera fulvicornis (Chrysomelidae), chrysomelid larvae and the bud-galling midge Dasineura rosaria did not differ between any of the plant categories. Hybrids were more severely infected by rust (Melampsora sp.) than S. caprea and the totally resistant S. repens. Densities of herbivores on hybrid willows were consistent with the dominance hypothesis (i.e. herbivore densities were similar to densities on one of the parental species) or supported the no-difference hypothesis. Furthermore, herbivore densities on hybrid plants were most similar to densities on the more susceptible parent. The breakdown in rust resistance in hybrid plants suggests that resistance traits are severely disrupted by the genetic re-arrangement in hybrids and that this increased susceptibility could select against hybridisation. Received: 17 February 1998 / Accepted: 15 June 1998     

Inheritance of Oryza sativa endornavirus in F1 and F2 hybrids between japonica and indica rice

We have found a 14 kbp double-stranded RNA (dsRNA) in many cultivars of japonica rice (Oryza sativa L.) but not in any cultivars of indica rice. This dsRNA is an RNA replicon with plasmid-like properties and is proposed to be a novel dsRNA virus, Oryza sativa endornavirus (OSV). Reciprocal crosses between the OSV-carrier japonica variety (Nipponbare) and the OSV-free indica variety (IR 26 or Kasalath) were performed to investigate whether OSV can be transmitted to F1 hybrids. When IR 26 and Nipponbare were used, efficient transmission of OSV from ova (93%) and pollen (89%) was observed. When Kasalath and Nipponbare were used, the OSV transmission efficiency to F1 progeny was 68% from ova and 20% from pollen. The transmission of OSV to F2 progeny plants was also complicated, showing non-Mendelian inheritance. These results suggest that the dsRNA replicon (OSV) is unstable in indica rice plants.     

Floral scent emission and pollinator attraction in two species of <Emphasis Type="Italic">Gymnadenia</Emphasis> (Orchidaceae)

We investigated scent composition and pollinator attraction in two closely related orchids, Gymnadenia conopsea (L.) R.Br. s.l. and Gymnadenia odoratissima (L.) Rich. in four populations during the day and night. We collected pollinators of both species using hand nets and sampled floral odour by headspace sorption. We analysed the samples by gas chromatography with mass spectrometry to identify compounds and with electroantennographic detection to identify compounds with physiological activity in pollinators. In order to evaluate the attractiveness of the physiologically active compounds, we carried out trapping experiments in the field with single active odour substances and mixtures thereof. By collecting insects from flowers, we caught eight pollinators of G. conopsea, which were members of four Lepidoptera families, and 37 pollinators of G. odoratissima, from five Lepidopteran families. There was no overlap in pollinator species caught from the two orchids using nets. In the scent analyses, we identified 45 volatiles in G. conopsea of which three (benzyl acetate, eugenol, benzyl benzoate) were physiologically active. In G. odoratissima, 44 volatiles were identified, of which seven were physiologically active (benzaldehyde, phenylacetaldehyde, benzyl acetate, 1-phenyl-2,3-butandione, phenylethyl acetate, eugenol, and one unknown compound). In field bioassays using a mixture of the active G. odoratissima compounds and phenylacetaldehyde alone we caught a total of 25 moths, some of which carried Gymnadenia pollinia. A blend of the active G. conopsea volatiles placed in the G. odoratissima population did not attract any pollinators. The two orchids emitted different odour bouquets during the day and night, but G. odoratissima showed greater temporal differences in odour composition, with phenylacetaldehyde showing a significant increase during the night. The species differed considerably in floral odour emission and this differentiation was stronger in the active than non-active compounds. This differentiation of the two species, especially in the emission of active compounds, appears to have evolved under selection for attraction of different suites of Lepidopteran pollinators.     

Influence of pollinator abundance and flower visitation on seed yield in red clover

Red clover (Trifolium pratense) is an important forage crop in temperate regions. However, the seed yield in red clover, especially in tetraploid cultivars, constrains the commercial success of this crop. The relationship between seed yield and several flower and inflorescence characteristics has been investigated previously, but up to now no clear link has been identified. The central questions in this article are as follows: (1) Whether nectar production and composition are correlated with seed yield? (2) Which is the main pollinator found on red clover in Flanders? (3) Whether pollinator preferences can explain differences between high and low seed-yielding cultivars and genotypes? We found that Bombus pascuorum was the most abundant pollinator on red clover in the region considered. Nectar volume, nectar composition or pollinator preference did not explain the variation in seed yield between genotypes, nor between diploid and tetraploid cultivars. To explain seed yield differences among red clover genotypes and ploidy levels better, we believe that plant-related characteristics like problems with meiosis, low pollen or seed viability or embryo abortion need to be investigated.     

Pollination ecology of Silene acutifolia (Caryophyllaceae): floral traits variation and pollinator attraction

BACKGROUND AND AIMS: The floral display influences the composition of pollinators interacting with a plant species. Geographic and temporal variation in pollinator composition complicates the understanding of the evolutionary consequences of floral display variation. This paper analyses the relationships between Silene acutifolia, a hermaphroditic perennial herb, and its pollinators, based on field studies in the north-west of Spain. METHODS: Studies were conducted over three years (1997-1999). Firstly, the main pollinators of this species were determined for two years in one population. Secondly, pollen limitation in fruit and seed production was analysed by supplementary hand pollinations, and counting the pollen grains and tubes growing in styles for two different-sized populations. Finally, the effect of flower size and number on the rate of visitation and total seed number was examined for 15 marked plants. RESULTS AND CONCLUSIONS: The primary pollinators were long-tongued insects, including Hymenoptera, Lepidoptera and Diptera, but the composition and visitation frequencies differed between years. Pollen limitation occurred in one of the years of study. There was between-population variation in the number of pollen grains and pollen tubes found in styles, suggesting pollen limitation in one population. Overall, pollinators visited plants with more open flowers more frequently, and pollinated more flowers within these plants. Conversely, petal and calyx sizes had no effect on insect visitation. Plants with higher rates of visits produced higher number of seeds, suggesting that pollinator-mediated limitation of seed and fruit production may be important in some years.     

Specialized pollination in the African milkweed Xysmalobium orbiculare: a key role for floral scent in the attraction of spider-hunting wasps

Specialized pollination by prey-hunting wasps is poorly documented in rewarding plants. Furthermore, the mechanisms of achieving specialization are not clear since flowers typically produce exposed nectar and have no morphological adaptations (such as long spurs) to exclude non-pollinating visitors. We investigated the pollination of Xysmalobium orbiculare and explored the functional roles of floral scent and nectar in attracting pollinators and deterring nectar robbers. Floral visitor observations showed that this milkweed is visited almost exclusively by pompilid wasps in the genus Hemipepsis. These wasps were the only insects to carry pollinia, and a cage experiment confirmed their effectiveness in removing and inserting pollinia on flowers. Hand-pollinations showed that plants are genetically self-incompatible and thus reliant on pollinators for seed set. Palatability experiments with honeybees showed that nectar is distasteful to non-pollinating insects and is therefore likely to play a functional role in deterring nectar thieves. Choice experiments in the field showed that the wasp pollinators are attracted primarily by floral scent rather than visual cues. Analysis of spectral reflectance of flowers revealed that flowers are dull colored and are unlikely to stand out from the background vegetation. We conclude that X. orbiculare is specialized for pollination by spider-hunting wasps in the genus Hemipepsis and utilizes floral scent to selectively attract its pollinators and unpalatable nectar to deter non-pollinating visitors.     

Cytology of F1 hybrids and chromosome number of F2 and BC1 progeny of the cross Bromus riparius x B. inermis

Summary Hybrids between B. inermis Leyss (2n=8x=56) and B. riparius Rehm. (2n=10x=70) were easily made. The F₁ hybrids had a fertility of 20%–50% under open pollination and backcrossing to B. inermis. Chromosome pairing in B. riparius was predominantly as bivalents (29.04–33.85 per cell for plant means). Bivalents also predominated in the F₁ hybrid (2n=9x=63) and there was a high level of pairing with no reduction in chiasma frequency. It was impossible to estimate the frequency of auto-versus allosyndetic pairing. Chromosome pairing in a hybrid between B. arvensis (2n=2x=14) and B. riparius confirmed that the B. riparius complement is capable of complete autosyndetic pairing. Chromosome numbers in the F₂ progeny ranged from 2n=56 to 72 but they were skewed towards 2n=63 to 70. Backcrosses ranged from 2n=56 to 63, as expected, with the distribution skewed towards 2n=56. Selection towards the 2n=56 level would be difficult in the F₂. Empirical observation suggested that cytoplasm had a major influence on morphology in the backcrosses. Additional studies are required to determine the best breeding scheme to introgress germ plasm between B. inermis and B. riparius.     

Vole preference for Salix caprea, S. repens, and their F1, F2, and backcross hybrids

In recent years, the influence of plant hybridization on plant–animal interactions has become an important issue; however, these studies have mainly focused on insects or slugs and to a large degree ignored mammalian herbivores despite their importance in many ecosystems. This study aims to determine the preference of voles for pure Salix caprea , S. repens , and their F1, F2, and backcross hybrids to evaluate whether voles select against hybrids in natural willow populations. To address this, we conducted two field studies and a cafeteria experiment in the laboratory with bank voles ( Clethrionomys glareolus ). We found no strong indications of reduced resistance in any of the hybrid taxa. Voles damaged more S. repens than S. caprea individuals. There was a general decreasing trend of utilization from pure S. repens to pure S. caprea , hybrid taxa being intermediate between the parents. Thus, voles seemingly do not have a more detrimental effect on hybrid fitness than on the fitness of pure individuals and probably will not select against hybrids in these willows populations.     

Effects of sex hormones and gender on attraction thresholds for volatile anal scent gland odors in ferrets

Body odors contribute to mate recognition and sexual partner preference in many mammals, including ferrets. We used a habituation/dishabituation procedure to test whether sex steroid hormones influence whether ferrets will approach and investigate different concentrations of volatile anal scent gland odors from male and female conspecifics. When tested with high concentrations of anal scent gland secretions in oil vehicle, gonadectomized male and female ferrets that received no sex steroids reliably discriminated anal scents from male and female conspecifics. This discrimination most likely reflects gender recognition rather than individual recognition because gonadectomized, sex steroid-treated ferrets discriminated between anal scents of males and females but not between anal scents of individual males or females. Treatment with either the estrogen receptor agonist, estradiol benzoate (EB), or the androgen receptor agonist, 5-alpha dihydrotestosterone proprionate (DHTP), increased investigation of low concentrations of anal scent by gonadectomized ferrets. These data suggest that ferrets could use anal scent gland secretions in mate recognition and that seasonal increases in circulating sex steroid hormones increase ferrets' responsiveness to low concentrations of these odors.     

Inheritance of flower color in periwinkle: orange-red corolla and white eye

The commonly found flower colors in periwinkle (Catharanthus roseus)--pink, white, red-eyed, and pale pink center--are reported to be governed by the epistatic interaction between four genes--A, R, W, and I. The mode of inheritance of an uncommon flower color, orange-red corolla and white eye, was studied by crossing an accession possessing this corolla color with a white flowered variety (Nirmal). The phenotype of the F(1) plants and segregation data of F(2) and backcross generations suggested the involvement of two more interacting and independently inherited genes, one (proposed symbol E) determining the presence or absence of red eye and another (proposed symbol O) determining orange-red corolla.     

Inheritance of gliadin proteins controlled by 1D chromosome in F2 hybrids oftriticum aestivum cultivars

The inheritance of the gliadin spectra in F₂ hybrids between cultivars hexaploid wheat Bezostaya 1 and Bisel has been investigated. The results from the electrophoretic examination and analysis of the gliadin spectra reveal that the components 1,2,3,11 and 15 from the Bezostaya 1 spectrum and 1,2, 3, 7,11 and 15 from the Bisel spectrum are controlled by allelic variants of one gliadin coding locus in 1D chromosome.