Genetic factors affecting food-plant specialization of an oligophagous seed predator.

Several ecological and genetic factors affect the diet specialization of insect herbivores. The evolution of specialization may be constrained by lack of genetic variation in herbivore performance on different food-plant species. By traditional view, trade-offs, that is, negative genetic correlations between the performance of the herbivores on different food-plant species favour the evolution of specialization. To investigate whether there is genetic variation or trade-offs in herbivore performance between different food plants that may influence specialization of the oligophagous seed-eating herbivore, Lygaeus equestris (Heteroptera), we conducted a feeding trial in laboratory using four food-plant species. Although L. equestris is specialized on Vincetoxicum hirundinaria (Apocynaceae) to some degree, it occasionally feeds on alternative food-plant species. We did not find significant negative genetic correlations between mortality, developmental time and adult biomass of L. equestris on the different food-plant species. We found genetic variation in mortality and developmental time of L. equestris on some of the food plants, but not in adult biomass. Our results suggest that trade-offs do not affect adaptation and specialization of L. equestris to current and novel food-plant species, but the lack of genetic variation may restrict food-plant utilization. As food-plant specialization of herbivores may have wide-ranging effects, for instance, on coevolving plant-herbivore interactions and speciation, it is essential to thoroughly understand the factors behind the specialization process. Our findings provide valuable information about the role of genetic factors in food-plant specialization of this oligophagous herbivore.

Genetic variation in herbivore resistance and tolerance: the role of plant life-history stage and type of damage.

Information of the patterns of genetic variation in plant resistance and tolerance against herbivores and genetic trade-offs between these two defence strategies is central for our understanding of the evolution of plant defence. We found genetic variation in resistance to two specialist herbivores and in tolerance to artificial damage but not to a specialist leaf herbivore in a long-lived perennial herb. Seedlings tended to have genetic variation in tolerance to artificial damage. Genetic variation in tolerance of adult plants to artificial damage was not consistent in time. Our results suggest that the level of genetic variation in tolerance and resistance depends on plant life-history stage, type of damage and timing of estimating the tolerance relative to the occurrence of the damage, which might reflect the pattern of selection imposed by herbivory. Furthermore, we found no trade-offs between resistance and tolerance, which suggests that the two defence strategies can evolve independently.

Host use of a hemiparasitic plant: no trade-offs in performance on different hosts.

To examine putative specialization of a hemiparasitic plant to the most beneficial host species, we studied genetic variation in performance and trade-offs between performance on different host species in the generalist hemiparasite, Rhinanthus serotinus. We grew 25 maternal half-sib families of the parasite on Agrostis capillaris and Trifolium pratense and without a host in a greenhouse. Biomass and number of flowers of the parasite were the highest when grown on T. pratense. There were significant interactions between host species and R. serotinus seed-family indicating that the differences in performance on the two hosts and without a host varied among the families. However, we found no significant negative correlations between performance of R. serotinus on the host species or between performance on the two hosts and autotrophic performance. Thus, the genetic factors studied here are not likely to affect the evolution of specialization of R. serotinus to the most beneficial host.

Relatedness affects competitive performance of a parasitic plant (Cuscuta europaea) in multiple infections.

Theoretical models predict that parasite relatedness affects the outcome of competition between parasites, and the evolution of parasite virulence. We examined whether parasite relatedness affects competition between parasitic plants (Cuscuta europaea) that share common host plants (Urtica dioica). We infected hosts with two parasitic plants that were either half-siblings or nonrelated. Relative size asymmetry between the competing parasites was significantly higher in the nonrelated infections compared to infections with siblings. This higher asymmetry was caused by the fact that the performance of some parasite genotypes decreased and that of others increased when grown in multiple infections with nonrelated parasites. This result agrees with the predictions of theories on the evolution of parasite virulence: to enhance parasite transmission, selection may favour reduced competition with genetically related parasites in hosts infected by several genotypes. However, in contrast to the most common predictions, nonrelated infections were not more virulent than the sibling infections.

Population structure of a parasitic plant and its perennial host.

Characterization of host and parasite population genetic structure and estimation of gene flow among populations are essential for the understanding of parasite local adaptation and coevolutionary interactions between hosts and parasites. We examined two aspects of population structure in a parasitic plant, the greater dodder (Cuscuta europaea) and its host plant, the stinging nettle (Urtica dioica), using allozyme data from 12 host and eight parasite populations. First, we examined whether hosts exposed to parasitism in the past contain higher levels of genetic variation. Second, we examined whether host and parasite populations differ in terms of population structure and if their population structures are correlated. There was no evidence that host populations differed in terms of gene diversity or heterozygosity according to their history of parasitism. Host populations were genetically more differentiated (F(ST) = 0.032) than parasite populations (F(ST) = 0.009). Based on these F(ST) values, gene flow was high for both host and parasite. Such high levels of gene flow could counteract selection for local adaptation of the parasite. We found no significant correlation between geographic and genetic distance (estimated as pairwise F(ST)), either for the host or for the parasite. Furthermore, host and parasite genetic distance matrices were uncorrelated, suggesting that sites with genetically similar host populations are unlikely to have genetically similar parasite populations.

Genetics of sex determination in the gynodioecious species Lobelia siphilitica: evidence from two populations.

In order to determine whether interactions between multiple sex-determining genes might be partly responsible for the wide variation in female frequency among populations of Lobelia siphilitica, we used progeny sex ratios from field-collected plants and from controlled crosses within and between two populations. We demonstrate that multiple cytoplasmic male-sterility types are present in a gynodioecious population where female frequency exceeds 50%. These male-sterility types each have corresponding nuclear alleles that can restore pollen fertility. Restoration of one male-sterility type appears to be controlled by a single, dominant allele, but restoration of a second cytoplasmic type is not easily explained with simple genetic models -- perhaps multiple nuclear loci and/or epistatic interactions are involved. In addition, the crosses show that pollen from some hermaphrodite plants in a population containing no females restores male fertility to plants from a geographically distant gynodioecious population that have male-sterile cytoplasm. Furthermore, some plants in that hermaphrodite population carry a male-sterile cytoplasm. Taken together, these results fit theoretical predictions that female frequency might be highly variable among populations when sex is determined by interactions between several nuclear and cytoplasmic genetic factors, some of which may not be present in all populations. The data also illustrate the need for more theoretical and empirical work investigating the evolutionary impact of nuclear restorer genes with complex action, and explaining the existence of nuclear restorers and cytoplasmic male-sterility genes in a population where females are very rare.

Local adaptation, resistance, and virulence in a hemiparasitic plant-host plant interaction.

Coevolution may lead to local adaptation of parasites to their sympatric hosts. Locally adapted parasites are, on average, more infectious to sympatric hosts than to allopatric hosts of the same species or their fitness on the sympatric hosts is superior to that on allopatric hosts. We tested local adaptation of a hemiparasitic plant, Rhinanthus serotinus (Scrophulariaceae), to its host plant, the grass Agrostis capillaris. Using a reciprocal cross-infection experiment, we exposed host plants from four sites to hemiparasites originating from the same four sites in a common environment. The parasites were equally able to establish haustorial connections to sympatric and allopatric hosts, and their performance was similar on both host types. Therefore, these results do not indicate local adaptation of the parasites to their sympatric hosts. However, the parasite populations differed in average biomass and number of flowers per plant and in their effect on host biomass. These results indicate that the virulence of the parasite varied among populations, suggesting genetic variation. Theoretical models suggest that local adaptation is likely to be detected if the host and the parasite have different evolutionary potentials, different migration rates, and the parasite is highly virulent. In the interaction between R. serotinus and A. capillaris all the theoretical prerequisites for local adaptation may not be fulfilled.

Phenotypic plasticity and priority rules for energy allocation in a freshwater clam: a field experiment.

We studied resource allocation among maintenance, reproduction and growth in the freshwater clam Anodonta piscinalis. Recent theoretical and empirical studies imply that organisms with indeterminate growth may have priority rules for energy allocation. That being so, the traits involved should potentially be capable of considerable phenotypic modulation, as a mechanism to adjust allocation. We tested this hypothesis using a 1-year reciprocal transplant experiment at six sites. Experimental clams were caged at higher than natural densities in order to detect any phenotypic modulation of the traits and discover the putative priority rules in energy allocation. We recorded the survival and shell growth of clams during the experiment, and the reproductive output, somatic mass and fat content of clams at the end of the experiment. Shell growth, somatic mass, and the reproductive output of females varied more among transplant sites than among the populations of origin, suggesting a high capacity for phenotypic modulation. However, the reproductive investment, somatic mass and shell growth were also affected by origin; clams from productive habitats invested more in reproduction and were heavier. In comparison to undisturbed clams, the reproductive output of the experimental clams was similar and their fat content was higher, whereas their shell growth was considerably slower and their somatic mass lower. These results suggests that when resources are limiting (due to high density) reproductive allocation overrides allocation to somatic growth. The highest mortality during the experiment coincided with the period of reproductive stress in the spring. Additionally, the proportion of reproducing females was lower in those transplant groups where the survival rate was lowest, suggesting that maintenance allocation overrides allocation to reproduction when available resources are scarce. The results of this field experiment support theoretical predictions and results of previous laboratory experiments that suggest that there are priority rules for energy allocation in organisms with indeterminate growth.

Effects of simulated herbivory on tillering and reproduction in an annual ryegrass,Lolium remotum.

This study examined the overall impact of simulated herbivory on tillering and reproductive performance of an annual ryegrass,Lolium remotum. The interaction between herbivore damage and intraspecific competition and the effect of the timing of damage were also studied. The experimental plants were sown at two densities and were randomly assigned to eight different damage treatments consisting of artificial leaf area removal by clipping with scissors or removal of one-third of the ripening seeds. The treatments were executed at two flowering stages. The pattern of tiller development differed significantly among treatments and between densities. At the lower density, earlier treatments delayed tiller development more than the same treatments executed later. At the higher density, all treatments delayed tiller development. The density effect was significant for all reproductive traits measured. The reproductive output of plants grown at the higher density was lower and the negative treatment effects were stronger than at the lower density. The treatment effect was significant for seed dry weight per plant and individual seed weight but not for number of seeds per plant. There were no statistically significant interaction effects between the damage treatments and density, suggesting that the plants responded to the damage similarly, irrespective of the density. The plants did not totally compensate for losses due to damage at either density, even though they slightly increased their resource allocation to sexual reproduction at the higher density.