Ectomycorrhizal colonization and intraspecific variation in growth responses of lodgepole pine

Across different host plant species, the effects of mycorrhizal colonization on host growth parameters can vary, but intraspecific variation in this relationship has rarely been measured. We tested the direction and consistency of the relationship between ectomycorrhizal colonization level and growth responses across seed families of Pinus contorta var. latifolia. Root tips of seedlings from eight full sib seed families varied in levels of ectomycorrhizal fungal colonization from 39% to 100%. We observed positive, negative, or neutral relationships between colonization level and shoot mass, depending on plant family. For the majority of seed families no relationship was observed between colonization level and root mass; however, two seed families showed negative relationships. Shoot height differed only by seed family. Results from our study indicate that the relationship between colonization level and host growth depends on host genotype. We suggest that models of plant intraspecific interactions should consider ectomycorrhizal associations when assessing phenotypic variability.     

Resistance and tolerance in a host plant-holoparasitic plant interaction: genetic variation and costs

Host organisms are believed to evolve defense mechanisms (i.e., resistance and/or tolerance) under selective pressures exerted by natural enemies. A prerequisite for the evolution of resistance and tolerance is the existence of genetic variation in these traits for natural selection to act. However, selection for resistance and/or tolerance may be constrained by negative genetic correlations with other traits that affect host fitness. We studied genetic variation in resistance and tolerance against parasitic infection and the potential fitness costs associated with these traits using a novel study system, namely the interaction between a flowering plant and a parasitic plant. In this system, parasitic infection has significant negative effects on host growth and reproduction and may thus act as a selective agent. We conducted a greenhouse experiment in which we grew host plants, Urtica dioica, that originated from a single natural population and represented 20 maternal families either uninfected or infected with the holoparasitic dodder, Cuscuta europaea. that originated from the same site. We calculated correlations among resistance, tolerance, and host performance to test for costs of resistance and tolerance. We measured resistance as parasite performance (quantitative resistance) and tolerance as the slopes of regressions relating the vegetative and reproductive biomass of host plants to damage level (measured as parasite biomass). We observed significant differences among host families in parasite resistance and in parasite tolerance in terms of reproductive biomass, a result that suggests genetic variation in these traits. Furthermore, we found differences in resistance and tolerance between female and male host plants. In addition, the correlations indicate costs of resistance in terms of host growth and reproduction and costs of tolerance in terms of host reproduction. Our results thus indicate that host tolerance and resistance can evolve as a response to infection by a parasitic plant and that costs of resistance and tolerance may be one factor maintaining genetic variation in these traits.     

Genetic Variation in Behavioral Response to Herbivore-Infested Plants in the Parasitic Wasp, Cotesia glomerata (L.) (Hymenoptera: Braconidae)

Female Cotesia glomerata (L.) relies on stimuli from herbivore-infested plants to select suitable hosts, but behavioral response to such stimuli is highly variable among individuals. This study investigates a genetic component of phenotypic variability in both short-range host-search and long-range host-location behaviors in the tritrophic system consisting of cabbage plants (Brassica oleracea L.), cabbage butterfly (Pieris brassicae L.) and the parasitoid, by comparing full-sib families established from a laboratory population and isofemale strains from a field population. Short-range host-search behaviors were examined within a Petri-dish test arena, and long-range host-location behaviors assessed in a wind tunnel. Significant differences among full-sib families were shown in the duration of walking on a plant-herbivore complex (i.e., a leaf section with two host caterpillars, their silk and feces) and searching off the complex, and the total time elapsed for wasps to locate a host larva after release into the test arena. Flight responses to and landing choices between the intact and the herbivore-infested plants were also significantly different among these families. Effects of families on both short-range host-search and long-range host-location behaviors were consistent, without significant influences of host larvae from which wasps emerged. The analysis of isofemale strains reveals that strains account for significant variation in the oriented flight response to herbivore-infested plants, and the isofemale heritability for this behavioral character is estimated as 0.447. The results suggest that genetic variation exists at different behavioral levels of the host-selection process in this parasitoid.     

Selection on offspring size varies within and among families in relation to host nutritional quality

Life history theory often assumes a positive relationship between offspring size and fitness, although the strength and form of this relationship is expected to vary with environmental conditions. In arthropods, surprisingly few studies have examined the influence of larval environment on the offspring size–fitness relation. In phytophagous insects, the few studies that have examined variation in larval host plants have found a negative correlation between host plant nutritional quality and the strength of selection favoring larger offspring size, suggesting that this pattern might be general. I present experimental evidence for such a relationship in a population of the moth Rothschildia lebeau feeding on its three primary host plant species. Unlike previous studies, I consider the effect of offspring size on growth and survival at two levels, both among families and among siblings within families. Neonate caterpillar mass had a significant effect on growth and survival. The effect on growth, however, was weak, resulted primarily from variation among families, and did not differ among host plant diets. The effect on survival was stronger and varied among host plant diets, among families, and within families on different host plants in a manner that was generally consistent with the hypothesized negative correlation between host plant nutritional quality and the strength of selection favoring larger offspring size. Overall, results suggest that the consequences of variation in offspring size for survival within and among families are host plant-dependent in this system.     

Phenotypic variation for adhesive tenacity in the barnacle Balanus amphitrite

Silicone fouling-release coatings represent a non-toxic alternative to biocide-containing ship hull paints. These coatings allow fouling organisms to attach to the hull surface, but prevent firm adhesion. Adhesive tenacity to fouling-release materials varies both among and within species. We quantified broad-sense genetic and environmental sources of intraspecific variation in tenacity to two silicone substrata, for the barnacle Balanus amphitrite. For both materials tenacity varied over an order of magnitude; however, the partitioning of this variation differed between the substrata. For International Veridian, a commercially-available fouling-release coating, removal stress varied significantly among maternal families and replicate barnacle cultures. Variation among the maternal families was associated with previously observed differences among these families in the condition of the adhesive plaque. Additional experiments suggested that variation among the replicate cultures arose from heterogeneity between replicate coatings in properties that affect tenacity. We could not attribute variation in removal stress for Dow Corning Silastic T-2, a silicone rubber used for mold-making, to any of the genetic or environmental sources tested. Instead, variation may have been due to measurement error or heterogeneity within replicate coatings in properties affecting tenacity. Differences among maternal families in removal stress may stem from variation in the interaction between the adhesive and the substratum, or in the viscoelastic properties of the adhesive plaque.     

Sequence variation and structural conservation allows development of novel function and immune evasion in parasite surface protein families

Trypanosoma and Plasmodium species are unicellular, eukaryotic pathogens that have evolved the capacity to survive and proliferate within a human host, causing sleeping sickness and malaria, respectively. They have very different survival strategies. African trypanosomes divide in blood and extracellular spaces, whereas Plasmodium species invade and proliferate within host cells. Interaction with host macromolecules is central to establishment and maintenance of an infection by both parasites. Proteins that mediate these interactions are under selection pressure to bind host ligands without compromising immune avoidance strategies. In both parasites, the expansion of genes encoding a small number of protein folds has established large protein families. This has permitted both diversification to form novel ligand binding sites and variation in sequence that contributes to avoidance of immune recognition. In this review we consider two such parasite surface protein families, one from each species. In each case, known structures demonstrate how extensive sequence variation around a conserved molecular architecture provides an adaptable protein scaffold that the parasites can mobilise to mediate interactions with their hosts.     

Host-associated fitness variation in a seed beetle (Coleoptera: Bruchidae): evidence for local adaptation to a poor quality host

The geographic distributions of many generalist herbivores differ from those of their host plants, such that they experience coarse-grained spatial variation in natural selection on characters influencing adaptation to host plants. Thus, populations differing in host use are expected to differ in their ability to survive and grow on these host plants. We examine host-associated variation in larval performance (survivorship, development time, and adult body weight) and oviposition preference, within and between two populations ofStator limbatus (Coleoptera: Bruchidae) that differ in the hosts available to them in nature. In one population,Acacia greggii (Fabaceae: Mimosoideae) andCercidium microphyllum (Fabaceae: Caesalpininoideae) are each abundant, while in the second population onlyC. floridum andC. microphyllum are present. In both populations, egg-to-adult survivorship was less than 50% onC. floridum, while survivorship was greater than 90% onA. greggii. Most of the mortality onC. floridum occurred as larvae were burrowing through the seed coat; very low mortality occurred during penetration of the seed coat ofA. greggii. Significant variation was present between populations, and among families (within populations), in survivorship and egg-to-adult development time onC. floridum; beetles restricted toCercidium in nature, without access toC. floridum, survived better and developed faster onC. floridum than beetles that had access toA. greggii. Large host effects on body size were detected for female offspring: females reared onA. greggii were larger than those reared onC. floridum, whereas male offspring wee approximately the same size regardless of rearing host. Trade-offs between performance onC. floridum andC. floridum were not detected in this experiment. Instead, our data indicate that development time and survivorship onC. floridum may be largely independent of development time and survivorship onA. greggii. Patterns of oviposition preference corresponded to the observed patterns of host suitability: in laboratory preference tests, beetles with access toA. greggii in nature tended to prefer this host more than beetles without access to this host in nature.     

Variation and covariation among life-history traits in Rumex acetosella from a successional old-field gradient

In this study morphological variation and the potential for competition to affect biomass and seedling selection of the families of five populations of Rumex acetosella L. sampled along a successional old-field gradient have been investigated. Seeds from 25 families were submitted to four competitive regimes: no competition (one plant per pot), medium competition (two plants/ pot taking plants from the same population), high within-population competition (four individuals from the same population in a pot) and high between-population competition (four individuals from two different populations in a pot). Eight traits were analysed after 3 months of growth for variation among families within populations. A significant difference among families within the two older populations was recorded for sexual biomass and related components. High sensitivity of these traits to density was observed in all populations except the youngest, suggesting specialization to particular environmental conditions in late successional populations, and a good adaptive capacity to buffer environmental variation in the pioneer population. Little significant interaction between competitive regimes and families within populations was found, i.e. genotypes within each population showed little variation in their response to environmental variation. Genotypic variance decreased with increasing competitive conditions for the majority of the traits. However, the percentage of variance in sexual reproduction explained by family was stable among treatments. Tradeoffs between vegetative reproduction and sexual reproduction were recorded at the population level along the successional gradient, with increasing competitive conditions. As succession proceeds, we observed a decrease in sexual reproduction and an increase in vegetative reproduction. At the family level, correlation among traits were similar when plants were grown in the absence of competition and at high density, with a significant negative correlation between sexual reproduction and vegetative reproduction. For both sprout number and sexual biomass, the performance of families grown under all the treatments was positively correlated. Together these results indicate allocational constraints on the reproductive biology of R. acetosella that may be favoured by natural selection and have influenced population differentiation along the successional gradient. However, they also revealed that the potential exists for evolutionary specialization through plasticity, in response to variation in environmental conditions.     

Genetic and phenotypic variation in diet breadth in a generalist herbivore

Summary The genetic and plastic components of polyphagy were investigated in a population ofLymantria dispar, the gypsy moth. A simple genetic experiment assessed the expression of (1) genetic variability in life history traits within each of four environments, (2) genetic variability in diet breadth, expressed as a change in the ranks of family performance across hosts, and (3) homeostasis (equivalent performance by a family across hosts) versus phenotypic plasticity (variable performance by a family across hosts). Sibs from each of 14 families, randomly selected from a single population, were reared on four diets: two natural hosts — chestnut and red oak, and two synthetic hosts — a standard laboratory diet and a low-protein version of this diet. Average population performance, measured in terms of development time and pupal weight, was better on standard laboratory diet than on low-protein diet, and was equal on chestnut and red oak for pupal weight, but better on chestnut oak for development time. Average population performance provided no information about the genetic component of host use ability. The gypsy moth expressed genetic variation in development time within each host environment and in pupal weight within natural host environments. Phenotypic plasticity was expressed by a significant number of families in development time and pupal weight across synthetic hosts and, to a lesser extent, across natural hosts. It was only across natural hosts that genetic variation in diet breadth was expressed, and this was confined to females. Genetic variability in diet breadth may be maintained in this species as a consequence of the unpredictability of its food sources.     

Resistance to 16 diverse species of herbivorous insects within a population of goldenrod,Solidago altissima: genetic variation and heritability

Summary Genetic variation in resistance to 16 species of herbivorous insects was studied in 18 clones of Solidago altissima growing in an old field near Ithaca, New York, USA. Resistance to each insect, defined as the abundance of a species attacking a particular host genotype relative to other genotypes, was measured in both the natural stand and in two experimental gardens. The heritability of resistance was estimated by parent-offspring regression and sibcorrelation. The primary result was that clones differed in resistance to 15 of 16 insect species. The resistance of genotypes to these insect species remained relatively constant over the four years of the study. However, for only 10 of these resistances were the heritability estimates significantly different from zero. Thus the common assumption of plant-insect studies — that phenotypic variation in insect abundance is closely correlated with underlying genetic variation — is only conditionally true. There is heritable variation in resistance to many insects, but not all. The insects for which we observed heritable variation in plant resistance represent five different orders and several functional groups, including leaf chewers, phloem and xylem feeders, and gall formers. There was no apparent pattern between the degree of heritability of plant resistance and the destructiveness, feeding method, breadth of host range, or taxonomic group of the insects. The lack of marked heritable variation in resistance to some insects may be the result of (a) reduced variation caused by strong selection during prolonged or repeated insect outbreaks, and (b) genotype-environment interactions that obscure differences among genotypes.     

Origins of P450 diversity

The P450 enzymes maintain a conserved P450 fold despite a considerable variation in sequence. The P450 family even includes proteins that lack the single conserved cysteine and are therefore no longer haem-thiolate proteins. The mechanisms of successive gene duplications leading to large families in plants and animals are well established. Comparisons of P450 CYP gene clusters in related species illustrate the rapid changes in CYPome sizes. Examples of CYP copy number variation with effects on fitness are emerging, and these provide an opportunity to study the proximal causes of duplication or pseudogenization. Birth and death models can explain the proliferation of CYP genes that is amply illustrated by the sequence of every new genome. Thus, the distribution of P450 diversity within the CYPome of plants and animals, a few families with many genes (P450 blooms) and many families with few genes, follows similar power laws in both groups. A closer look at some families with few genes shows that these, often single member families, are not stable during evolution. The enzymatic prowess of P450 may predispose them to switch back and forth between metabolism of critical structural or signal molecules and metabolism dedicated to environmental response.     

Variant antigen gene expression in malaria

Pathogens of the genus Plasmodium are unicellular parasites that infect a variety of animals, including reptiles, birds and mammals. All Plasmodium species target host erythrocytes and replicate asexually within this niche. In humans, proliferation within erythrocytes causes disease symptoms ranging from asymtomatic infection to severe disease, including mild to severe febrile and respiratory symptoms, profound anaemia and obstruction of blood flow. The most serious form of human malaria is caused by Plasmodium falciparum, a pathogen that is responsible for several million deaths annually throughout the developing world. Malaria parasites succeed in evading the host immune response to establish long-term, persistent infections, thus increasing the efficiency by which they are transmitted to the mosquito vector. The ability to evade the host immune system, in particular the avoidance of antibody-mediated immunity against parasite-encoded surface proteins, is the result of amplification of extensive repertoires of multicopy, hypervariable gene families that encode infected erythrocyte or merozoite surface proteins. Via switching between antigenically diverse genes within these large families, populations of parasites have the capacity for rapid variation in antigenicity and virulence over the course of an infection. Here we review the amplification and generation of antigenic diversity within the Plasmodium variant gene families, as well as discuss the mechanisms underlying their tightly controlled gene expression and antigenic switching.     

Host–parasite coevolution: genetic variation in a virus population and the interaction with a host gene

Host–parasite coevolution is considered to be an important factor in maintaining genetic variation in resistance to pathogens. Drosophila melanogaster is naturally infected by the sigma virus, a vertically transmitted and host‐specific pathogen. In fly populations, there is a large amount of genetic variation in the transmission rate from parent to offspring, much of which is caused by major‐effect resistance polymorphisms. We have found that there are similarly high levels of genetic variation in the rate of paternal transmission among 95 different isolates of the virus as in the host. However, when we examined a transmission‐blocking gene in the host, we found that it was effective across virus isolates. Therefore, the high levels of genetic variation observed in this system do not appear to be maintained because of coevolution resulting from interactions between this host gene and parasite genes.     

DNA barcoding of endoparasitoid wasps in the genus Anicetus reveals high levels of host specificity (Hymenoptera: Encyrtidae)

The genus Anicetus includes economically important biocontrol agents that are introduced for control of soft and wax scale insect agricultural pests (Ceroplastes spp.). Understanding of host–parasitoid associations is critical to the successful outcome of their utilization in biological control projects. However, identification of these parasitoids is often difficult because of their small size and generally similar morphological features, and hence, studies on the host–parasitoid associations. Here, nucleotide sequence data were generated from the mitochondrial COI gene and the D2 region of 28S rRNA to assess genetic variation within and between species of Anicetus occurring in China. The results of this study support the use of the COI and the D2 region of 28S rRNA gene as useful markers in separating species of Anicetus, even in cases where morphological differences are subtle. On the other hand, the COI gene is also useful in recognizing species with much variation in morphology. DNA barcoding reveals high levels of host specificity of endoparasitoids wasps in the genus Anicetus. Our results indicate that each Anicetus species is adapted to a limited set of host species, or even are monospecific in their host choice.     

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.     

Liriomyza huidobrensis in Yunnan, China: current distribution and genetic structure of a recently established population

Liriomyza huidobrensis Blanchard (Diptera: Agromyzidae) is a very serious and economically important pest around the world. Liriomyza huidobrensis in China was first reported from Kunming of Yunnan province in 1993. We report here that this pest has recently expanded its distribution, along with a host plant range extension and population explosion. The mitochondrial cytochrome oxidase II gene was sequenced for eight populations from Yunnan. All individuals were identical: no genetic variation was observed between populations either from different geographical localities or from different host plants. The phylogenetic analysis shows that the Yunnan population is grouped into the South American clade, which also includes other recently introduced Asian populations. Together with ecological data and colonization history of this pest, our results suggest that Yunnan population might have an ultimate, albeit not immediate, origin from South American populations.