The relative importance of architecture and resource competition in allocation to pollen and ovule number within inflorescences of Hosta ventricosa varies with the resource pools

Background and Aims

Allocation of resources to floral traits often declines distally within inflorescences in flowering plants. Architecture and resource competition have been proposed as underlying mechanisms. The aim of the present study is to assess the relative importance of resource competition and architectural effects in pollen and ovule production on racemes of Hosta ventricosa, an apomictic perennial herb.

Methods

Combinations of two defoliation treatments (intact and defoliated) and two fruit-set treatments (no-fruit and fruit) were created, and the roles of architecture and resource competition at each resource level were assessed.

Key Results

Pollen and ovule number per flower increased after defoliation, but pollen to ovule ratio per flower did not change. Pollen, ovules and the pollen to ovule ratio per flower declined distally on racemes at each resource level. In the intact treatment, fruit development of early flowers did not affect either pollen or ovule number of late flowers. In the defoliated treatment, fruit development of early flowers reduced both pollen and ovule numbers of late flowers due to over-compensation caused by defoliation. Late flowers on defoliated fruit racemes produced less pollen than intact fruit racemes but the same number of ovules; therefore, the reduction in pollen number was not caused by over-compensation. In addition, the fruit-set rate of early flowers during flowering was higher in intact racemes than in defoliated racemes.

Conclusions

In flowering plants, the relative importance of architecture and resource competition in allocation to pollen and ovules may vary with the resource pools or the overall resource availability of maternal plants.     

Seed-coat anatomy of japanese species ofCorydalis andDicentra (Papaveraceae; Fumarioideae)

Development and structure of seed-coat were examined in 16 species ofCorydalis and two species ofDicentra. Neither the tegmen (developed ii) nor the testa (developed oi) is multiplicative during seed development. Mature seed-coat is consistent in all the species examined in having mechanical structure in the exo- or endotesta. Differences are found in (1) which of the integumentary layers develops into the main mechanical layer (exotesta or endotesta), (2) whether a mesotesta is differentiated or not, and (3) whether the endotegmen is persistent as a layer of thick-walled cells, is persistent only partially, or is thoroughly degenerated. Theses seed-coat characters distinguish six groups of species (i.e., four groups inCorydalis and two inDicentra), which represent well infrageneric taxa proposed on the other characters. Evidence from seed-coat anatomy further suggests thatDicentra spectabilis (subg.Hedycapnos) retains the primitive (endotestal) seed-coat of Fumarioideae.     

Pollen size-number trade-off and pollen-pistil relationships in Pedicularis (Orobanchaceae)

Current patterns of floral design in Pedicularis must have undergone an evolutionary process of interacting among components of floral traits, and then formed internal relationships among these traits. To detect such correlations, which may provide insight to understand flower evolution, 40 Pedicularis species representing all corolla types of the genus were studied. Results show that, interspecifically, pollen size correlates negatively with pollen number, but positively with pistil length. This suggests that plants evolve an optimal pollen size, which balances the advantages of large pollen size for gametophytic competition against the fecundity disadvantages of fewer pollen grains. In contrast to sex allocation theory, this study does not find a trade-off, but an interspecific positive correlation between pollen and ovule number. This is consistent with the hypothesis that genetic variation for resource acquisition may in part be responsible for the lack of negative correlation between male and female function.This work was supported by the State Key Basic Research and Development Plan, China (Grant No. G2000046804) to YHG. The authors would like to thank Peter K. Endress and two anonymous reviewers for providing critical comments and helpful suggestions, Qing-Feng Wang, Jing-Yuan Wang and Jin-Ming Chen for their helpful suggestions. Shi-Guo Sun, Jing Xia, and Qian Yu are thanked for their assistance in both the field work and laboratory phases of the project.     

Infrageneric variability of pollen morphology in Palaua (Malveae,Malvaceae) and the taxonomic utility of quantitative pollen characters

Palaua (Malvaceae) comprises 15 species endemic to the coastal deserts of Chile and Peru. Previous molecular phylogenetic analyses have shown that this genus is monophyletic and can be subdivided into three clades. In the present study, pollen morphology of all species of Palaua was examined using light and scanning electron microscopy to determine whether it provides additional data in support of the proposed infrageneric groups. The pollen grains are spheroidal, medium to large in diameter, spinose and pantocolporate – the ectocolpi are very short. The tectum is perforate and characterised by granula. In all species the nexine is of a similar thickness in all areas of the pollen grain, while the sexine thickness varies, being notably thicker in the areas where the broad spine bases are sited (‘spine cushions’). Many of the quantitative characters have conspicuous variability, but the variability shows considerable overlap between species. Nonetheless, P. guentheri, P. inconspicua, P. malvifolia and P. modesta are united as a group by having the smallest pollen grains with the smallest apertures, the shortest spines and the shortest interspinal distance. This grouping reflects only partially the suggested infrageneric clades, although it does tend to unite the species with the smallest flowers. A possible link between reproductive resources and pollen size is also considered, as well as the influence of polyploidy. However, the taxonomic utility of these quantitative characters is weakened by species with more than one cytotype.     

Variation in structural gender in the hermaphrodite Helleborus foetidus (Ranunculaceae): within- and among-population patterns

In hermaphrodite plants, variations in structural gender (defined as the ratio between male and female gametes) may occur at different levels (among flowers, plants, and populations). In this study, we investigated variation in four traits influencing structural gender (number of carpels, ovules per carpel, stamens, and pollen grains per stamen) within and among six distant populations of the hermaphrodite perennial herb Helleborus foetidus (Ranunculaceae) in the Iberian Peninsula. Our results show that the four traits investigated varied significantly at all levels considered. Traits influencing the female sexual component (number of carpels and ovules per carpel) showed greater variation at the lowermost levels (within flower and plant) than traits influencing the male component, which in turn varied more markedly among populations. Number of carpels per flower and number of pollen grains per anther were the most important traits affecting between-plant variation in structural gender. There was no evidence of significant plant-level trade-offs or correlations between the various male and female traits, which covaried differently across populations. The observed between-population variation in structural gender of Helleborus foetidus can be explained as a consequence of differences in self-pollination levels related to a flowers mating environment.     

Temporal variation in the pollen:ovule ratios of Clarkia (Onagraceae) taxa with contrasting mating systems: field populations

Among plants, pairs of selfing vs. outcrossing sister taxa provide interesting systems in which to test predictions concerning the magnitude and direction of temporal changes in sex allocation. Although resource availability typically declines towards the end of the growing season for annual taxa, temporal changes in mating opportunities depend on mating system and should change less in selfing taxa. Consequently, given that the pollen:ovule (P:O) ratio of flowers reflects the investment in (and potential fitness pay-off due to) male vs. female function, we predicted that the P:O ratio should also be less variable among and within selfers than in closely related outcrossers. To test these predictions, we measured temporal changes in sex allocation in multiple field populations of two pairs of sister taxa in the annual flowering plant genus Clarkia (Onagraceae). In the outcrossing Clarkia unguiculata and the selfing Clarkia exilis, ovule production declined similarly from early to late buds, whereas pollen production remained constant or increased in the outcrosser but remained constant or decreased in the selfer. Consequently, the P:O ratio increased within unguiculata populations but marginally increased or stayed constant in exilis populations. In all populations of the selfing Clarkia xantiana spp. parviflora and the outcrossing C. x. spp. xantiana, both ovule and pollen production per flower declined over time. The effects of these declines on the P:O ratio, however, differed between subspecies. In each xantiana population, the mean P:O ratio did not differ between early and late flowers, although individuals varied greatly in the direction and magnitude of phenotypic change. By contrast, parviflora populations differed in the mean direction of temporal change in the P:O ratio. We found little evidence to support our initial predictions that the P:O ratio of the selfing taxa will consistently vary less than in outcrossing taxa.     

Variation and geographical distribution of ploidy levels inPennisetum sectionBrevivalvula (Poaceae) in Burkina Faso,Benin and southern Niger

Pennisetum sect.Brevivalvula is a species complex characterized by polyploidy and apomixis. Ploidy level was assessed by DAPI-flow cytometry for 304 plants of the section, originating from Burkina Faso, Benin and southern Niger. The results were confirmed for 54 plants based on chromosome counts. The samples show four euploidy levels (with x = 9) distributed among five species:P. hordeoides (2n = 36, 54),P. pedicellatum (2n = 36, 45, 54),P. polystachion (2n = 18, 36, 45, 54),P. setosum (2n = 54), andP. subangustum (2n = 18, 36, 54). The geographical distribution of these ploidy levels seems related to major vegetation zones present in Africa. Diploid populations ofP. polystachion andP. subangustum were found in the Banfora area, in Burkina Faso.     

Factors affecting intrafruit pattern of ovule abortion and seed production in Hormathophylla spinosa (Cruciferae)

We investigated the pattern of intrafruit seed production in Hormathophylla spinosa (Cruciferae) over a 7-yr period. H. spinosa fruit have two chambers, each chamber containing two ovules, and usually situate perpendicular to the infructescence axis with one chamber above the other. The percentage of ovules ripening to seeds is usually lower than 50%. In addition, we found a consistent position-dependence in seed ripening probability, since more than 90% of the ripe seeds are arranged in the lower chamber of the fruit. This pattern of seed production is not produced by the direct or indirect effect of seed predators, by pollen limitation or by nonrandom fertilizations. By contrast, fruit removal experiments showed that sibling rivalry can partially explain the seed production pattern, be the cause of the reduction in the seed number from the uppermost threshold of 50%. Moreover, experimental manipulations of fruit orientation suggest that the position-dependence in seed production is the cause but not the effect of seed ripening. We presume that some architectural effect is producing a significant increase in the ripening probability of the ovules arranged in the lower chamber, causing thus the observed pattern in the intrafruit seed production in H. spinosa and severely constraining the maximum number of seeds per fruit to two.     

Contribution to the taxonomy and phylogeny of Sarcocapnos DC. (Fumariaceae)

 To solve problems concerning the status of the taxa described in the genus Sarcocapnos, we have conducted a study using morphological, pollen morphology (light microscopy), cytogenetic and molecular techniques. Focusing on the last technique, we have sequenced ITS-1 and ITS-2 of nuclear rDNA. The species differ basically according to 5 morphological traits (leaf shape, flower spur, corolla colour, corolla size, and crest of the stigmatic surface). The cytogenetic analyses indicated n=16 to be the standard chromosome number. The ITS analyses showed that the genus is monophyletic, defining two main well-supported clades, one containing S. saetabensis and S. enneaphylla, and one containing the rest of the species. In this second clade, S. speciosa, S. pulcherrima, and S. baetica subsp. ardalii are related, as are S. integrifolia, S. crassifolia subsp. crassifolia, and S. crassifolia subsp. atlantis; S. baetica subsp. baetica forms a trichotomy with the foregoing groups. S. speciosa is shown to be a species separate from S. crassifolia subsp. crassifolia, as in the case of S. baetica with respect to S. integrifolia. Palynologically, the parameters used enabled us to establish clear differences between the taxa, often corroborating the macromorphological and genetic data. The flower spur has been reduced several times in different groups of the genus, for which the classifications established on the basis of this trait are paraphyletic. Received July 16, 2002; accepted December 11, 2002 Published online: March 31, 2003     

A phenetic comparison of someFumaria spp. (Fumariaceae)

The taxonomy of the genusFumaria has not been considered in detail sincePugsley's work in 1919 ff., and few modern methods have been applied to it. In a phenetic study, 33 populations of 11Fumaria spp. were grown in uniform conditions, and seven morphological characters measured. After re-expression and transformation the data were analysed by cluster analysis and principal components analysis. Alternative analyses did not indicate contradictory taxonomic conclusions. Artificial crosses gave some evidence on interfertility, and suggestedF. occidentalis to be an allopolyploid ofF. bastardii ×F. capreolata. Pugsley's subsectional classification is supported within sect.Parviflora, but not within sect.Grandiflora. His two sections are seen to be meaningful, but not discrete.     

The size and germinability of Scots pine pollen in different temperatures in vitro

The effect of genotype, the origin of genotype, and germination temperature on Scots pine pollen grain size, hydration rate, germinability, and tube growth was studied in vitro. The mean sizes of dry and germinated pollen grains varied among pollen genotypes in different ways, thus the hydration rate varied among genotypes. Pollen from Scots pine that originates in northern Finland hydrated more than pollen from a population in southern Finland. Germination temperature had no effect on the hydration rate. Germinability and tube growth rate of northern genotypes were higher at 20 °C than at 15 °C. Differences among southern genotypes were not significant. At 15 °C, the germinability and pollen tube growth rate of northern genotypes were lower than southern genotypes. At 20 °C, the differences were not significant. It appears that germination and growth of pollen from northern populations are enhanced at higher temperatures whereas pollen from southern populations is unaffected.     

Natural levels of pollination intensity and effects of pollen loads on offspring quality in females of Thalictrum pubescens (Ranunculaceae)

Females of Thalictrum pubescens produce stamens containing sterile pollen. Earlier studies have shown the presence of stamens does not increase seed set through increased visitation by insects. Insects may, however, increase pollen deposition on stigmas and increase pollen competition. This paper examines: 1) pollen deposition levels in natural populations and 2) whether larger pollen loads lead to higher quality offspring. The majority of stigmas in two populations received less than 15 grains, but a small proportion had large loads. The latter may represent the occasional insect visit. These visits may provide the opportunity for pollen tube competition. In the greenhouse, flowers receiving heavy-pollen loads had higher seed set than flowers receiving light-pollen loads. Seeds from the two treatments were the same mass, had similar germination times, and seedlings had the same mass. In conclusion, it does not appear that pollen tube competition can account for the maintenance of stamens in females of T. pubescens.     

Distyly and pollen dimorphism inLinum suffruticosum (Linaceae)

The Mediterranean distylousLinum suffruticosum has dimorphic pollen. That from short-styled plants has sub-monomorphic surface excrescences with minute papillae; that from long-styled plants has strongly dimorphic excrescences with conspicuous papillae. The differences are much like those described in species of other sections of the genus. Pollen of the related homostylousL. tenuifolium has monomorphic pollen. It combines characteristics of bothL. suffruticosum types but resembles that of the long-styled plants somewhat more closely. Ribbonlike styles of long-styled plants ofL. suffruticosum provide another feature which seems to be controlled by the S supergene thought to be associated with distylic incompatibility. The pollen ofL. suffruticosum andL. tenuifolium is very similar to that of some species of sectionLinastrum and would support their placement in that section.     

The classification and evolution of pollen types ofPlanchonioideae (Lecythidaceae)

The pollen type ofPlanchonioideae is unique among the angiosperms in its syntricolpatism associated with the marginal ridge, marginal groove, and polar cushion; it has been studied byJ. Muller, who classified it into three types, mainly with a light microscope. In the present study, pollen grains of the six constitutent genera were studied with a scanning electron microscope, five additional pollen types were recognized. Of the eight pollen types the one with sealed polar ectoapertures and weakly developed marginal ridges, without marginal grooves and polar cushions is regarded as the most primitive. Evolutionary trends proceeded with the opening of the polar ectoapertures, the complication of the polar cushions and marginal grooves. In the most advanced form, the marginal ridges and marginal grooves coalesced to form two concentric rings, whereas the polar cushions were secondarily reduced.     

Temporal patterns of ovule allocation,fruit set,and seed predation inAnemonopsis macrophylla (Ranunculaceae)

Ovule deployment patterns were studied inAnemonopsis macrophylla Sieb. & Zucc. to develop hypotheses about major selective forces underlying different temporal strategies. Carpel number per flower decreased significantly with relative flowering date. The trend was present at individual, population, and among-population, levels. Both ovule number per carpel and ripe seed weight decreased with later flowering date. Such temporal allocation patterns may be adaptive in late-flowering species with extended flowering patterns, because successively later flowers run a higher risk of lower resource availability and premature death due to deteriorating climatic conditions.     

Reproductive isolation and hybridization in sympatric populations of three Dactylorhiza species (Orchidaceae) with different ploidy levels

Background and Aims

The potential for gene exchange between species with different ploidy levels has long been recognized, but only a few studies have tested this hypothesis in situ and most of them focused on not more than two co-occurring species. In this study, we examined hybridization patterns in two sites containing three species of the genus Dactylorhiza (diploid D. incarnata and D. fuchsii and their allotetraploid derivative D. praetermissa).

Methods

To compare the strength of reproductive barriers between diploid species, and between diploid and tetraploid species, crossing experiments were combined with morphometric and molecular analyses using amplified fragment length polymorphism markers, whereas flow cytometric analyses were used to verify the hybrid origin of putative hybrids.

Key Results

In both sites, extensive hybridization was observed, indicating that gene flow between species is possible within the investigated populations. Bayesian assignment analyses indicated that the majority of hybrids were F₁ hybrids, but in some cases triple hybrids (hybrids with three species as parents) were observed, suggesting secondary gene flow. Crossing experiments showed that only crosses between pure species yielded a high percentage of viable seeds. When hybrids were involved as either pollen-receptor or pollen-donor, almost no viable seeds were formed, indicating strong post-zygotic reproductive isolation and high sterility.

Conclusions

Strong post-mating reproductive barriers prevent local breakdown of species boundaries in Dactylorhiza despite frequent hybridization between parental species. However, the presence of triple hybrids indicates that in some cases hybridization may extend the F₁ generation.     

Flow cytometric measurements do not reveal different ploidy levels in Minthostachys (Lamiaceae)

Ten of the 17 species of the taxonomically difficult Andean mint genus Minthostachys (Lamiaceae) were submitted to flow cytometric measurements of nuclear DNA content to test the hypothesis of the occurrence of different ploidy levels within the genus. Nuclear DNA content was found to vary from 1.643 to 1.775 pg, i.e by only ca. 8% between individual accessions, thus providing no evidence for polyploidy in Minthostachys. While these results do not preclude the possibility that the genus contains polyploid species nor the occurrence of heteroploidy with nearly identical nuclear DNA contents, they suggest that polyploidy did not play a major role in its diversification.     

Variability of microsatellites BM224 and Bcal7 in populations of green toads (Bufo viridis complex) differing by nuclear DNA content and ploidy

The variability of microsatellites BM224 and Bcal7 was studied for the first time in three species of the diploid-polyploid complex of Bufo viridis (B. viridis, B. oblongus, and B. pewzowi). The locus Bcal7 was established to be monomorphic in all samples studied. In microsatellite BM224, three allele variants were found. Among tetraploid toads, the western Asiatic species B. oblongus was characterized by one allele only, the eastern B. pewzowi, by the two other alleles. A similar distribution was also revealed in triploid individuals on the borders of range between tetraploid and diploid species. Among the diploid species B. viridis samples, all three allele variants of microsatellite BM224 were observed. Their distribution in the area proved to be geographically determined. In diploid toads, a similarity was revealed between the distribution of microsatellite BM224 alleles and variability of the nuclear DNA content.     

Fluctuations of grass pollen content in the atmosphere of East Perugia and meteorological correlations (year 1989)

Summary Gramineae pollination from a pollen monitoring station located in the eastern suburb of Perugia and meteorological correlations are reported. The data refers to the year 1989. Grass pollen peak pollination was from May to July; in this period the influence of relative humidity and of temperature on pollen concentration was very high. Phenological observations, to identify the time of maximum stamen extension in the most common genera in the area, are also reported. During the period of investigation the counts of pollen grains over four-hour periods showed a regular diurnal rhythm with peaks of concentration in the four-hour period 16.00–20.00. Aerosporological data and meteorological data related to four-hour periods were correlated following different criteria.