TWO HERBIVORES AND CONSTRAINTS ON SELECTION FOR RESISTANCE IN BRASSICA RAPA

Although most plants experience herbivory by several insect species, there has been little empirical work directed toward understanding plant responses to these simultaneous selection pressures. In an experiment in which herbivory by flea beetles (Phyllotreta cruciferae) and diamondback moths (Plutella xylostella) was manipulated in a factorial design, I found that selection for resistance to these herbivores is not independent in Brassica rapa. Specifically, the effect of flea beetle damage on B. rapa fitness depends on the amount of diamondback moth damage a plant experiences: damage by these herbivores has a nonadditive effect on plant fitness. When diamondbacks are abundant, plants that sustain high levels of damage by flea beetles are favored by natural selection, but when diamondbacks are rare, a low level of damage by flea beetles is favored. However, resistance to the later-feeding diamondback moth is not affected by the presence or absence of damage by early-feeding flea beetles. Thus, there are no plant-mediated ecological interactions between these herbivores that affect the outcome of selection for resistance. Because these herbivores do not independently affect plant fitness, neither is likely to develop a pairwise coevolutionary relationship with its host. Instead, coevolution is diffuse.     

Consequences of sequential attack for resistance to herbivores when plants have specific induced responses

Plants in nature are attacked sequentially by herbivores, and theory predicts that herbivore-specific responses allow plants to tailor their defenses. We present a novel field test of this hypothesis, and find that specific responses of Solanum dulcamara lead to season-long consequences for two naturally colonizing herbivores, irrespective of the second herbivore to attack plants. This result indicates that responses induced by the initial herbivore made plants less responsive to subsequent attack. We show that initial herbivory by flea beetles and tortoise beetles induce distinct plant chemical responses. Initial herbivory by flea beetles lowered the occurrence of conspecifics and tortoise beetles relative to controls. Conversely, initial herbivory by tortoise beetles did not influence future herbivory. Remarkably, the experimentally imposed second herbivore to feed on plants did not modify consequences (induced resistance or lack thereof) of the first attacker. Induction of plant chemical responses was consistent with these ecological effects; i.e. the second herbivore did not modify the plant's initial induced response. Thus, canalization of the plant resistance phenotype may constrain defensive responses in a rapidly changing environment.     

Herbivore-induced responses and patch heterogeneity affect abundance of arthropods on plants

Abstract.  1. Plants respond to herbivore damage by inducing defences that can affect the abundance of herbivores and predators. These tritrophic interactions may be influenced by heterogeneity in plant neighbourhood.
2. In the present study, the effects of induced responses on the abundance of herbivores (flea beetles and aphids), omnivores (pirate bugs and thrips), and predators (lady beetles and spiders) on individual plants and their neighbours between and within patches composed of three tomato plants was investigated.
3. Herbivore damage was manipulated to create homogeneous patches where either all or none of the plants had defences induced by herbivore damage, and heterogeneous patches where only one of the plants was induced.
4. Arthropod abundance on plants at different scales was compared by testing between patch effects (patch level), for neighbourhood effects at the plant phenotype level (neighbourhood level), and between near and far plants (within patch position).
5. At the patch level , plants in homogeneously induced patches contained fewer flea beetles and pirate bugs, but more lady beetles, compared with homogeneously non-induced patches. There was no effect of patch type on the abundance of aphids, thrips, and spiders on plants.
6. At the neighbourhood level , induced plants in heterogeneous patches contained more flea beetles and pirate bugs compared with induced plants in homogeneous patches, indicating that the abundance of some herbivores and omnivores on induced plants varied depending on the phenotype of the other plants within the patch. Within patch position, there was no evidence that the abundance of herbivores or predators on non-induced plants was affected by proximity to an induced plant.
7. Therefore, variation in plant neighbourhood generated by induced plant responses affected the abundance of three arthropods from three feeding guilds.     

Negative interactions between chemical resistance and predators affect fitness in soybeans

Abstract 1. Plants may benefit from both chemical resistance traits and the presence of predators of herbivores. In past studies, the interaction between resistance and predators varies from complementary to antagonistic among different systems. However, this interaction has primarily been quantified by effects on predator abundance or vigor, not effects on plant fitness. 2. In this study, the combined effects of chemical resistance and predators on plant fitness were examined using soybeans (Glycine max), herbivorous Mexican bean beetles (Epilachna varivestis), and predaceous spined soldier bugs (Podisus maculiventris). Mexican bean beetles were reared in field cages in the presence or absence of spined soldier bugs on soybeans with or without strong constitutive chemical resistance. 3. Spined soldier bugs were more likely to feed on Mexican bean beetles that fed on susceptible than on resistant plants. 4. Susceptible plants with predators produced significantly more seeds than those without predators, while resistant plants did not produce significantly different numbers of seeds based on the presence or absence of predators. 5. Selection for the production of some types of chemical resistance in plants would thus be expected to be stronger with lower predation rates. 6. These results also suggest predator introductions would be more effective on plants without a strong constitutive chemical resistance to herbivores.     

EVOLUTIONARY ECOLOGY OF DATURA STRAMONIUM L. IN CENTRAL MEXICO: NATURAL SELECTION FOR RESISTANCE TO HERBIVOROUS INSECTS

It has been assumed that herbivores constitute a selective agent for the evolution of plant resistance. However, few studies have tested this hypothesis. In this study, we look at the annual weed Datura stramonium for evidence of current natural selection for resistance to herbivorous insects. Paternal half-sib families obtained through controlled crosses were exposed to herbivores under natural conditions. The plants were damaged by two folivorous insects: the tobacco flea beetle Epitrix parvula and the grasshopper Sphenarium purpurascens. Selection was estimated using a multiple-regression analysis of plant size and of damage by the two herbivores on plant fitness measured as fruit production for both individual phenotypes and family breeding values (genetic analysis). Directional phenotypic selection was detected for both larger plant size and lower resistance to the flea beetles, whereas stabilizing phenotypic selection was revealed for resistance to S. purpurascens. However, performing the same analyses on the breeding values of the characters revealed directional and stabilizing selection only for plant size. Thus, no agreement existed between the results of the two types of analyses, nor was there any detectable potential for genetic change in the studied population because of selection on herbivore resistance. The narrow-sense heritability of every trait studied was small (all <0.1) and not different from zero. The potential for evolutionary response to natural selection for higher resistance to herbivores in the studied population of D. stramonium is probably limited by lack of genetic variation. Natural selection acts on phenotypes, and the detection of phenotypic selection on resistance to herbivores confirms their ecological importance in determining plant fitness. However, evolutionary inferences based solely on phenotypic selection analyses must be interpreted with caution.     

Competition among herbivores of Solanum carolinense as a constraint on the evolution of host-plant resistance

Interspecific competition (or facilitation) between herbivores sharing a host plant species can result in negative (or positive) correlations in damage levels, independent of a plant’s genetic covariance in resistance to the herbivores. Just like genetic correlations in resistance to herbivory, these “environmental correlations” in damage may affect the evolution of resistance in the host plant. In a field study of 960 ramets and 40 genets of the herbaceous plant Solanum carolinense, I looked for evidence of such environmental correlations in damage caused by 11 species of herbivores, including 10 insects and 1 mammal. There were 28 significant correlations in damage levels between species (21 negative and 7 positive) after plant genetic influences on resistance were statistically removed. Negative environmental correlations were more likely between species that fed upon the same type of plant organs than between those that fed on different types of organs, and the magnitudes of the correlations were inversely proportional to the abundance of the organ types. Taken together, these results offer strong evidence that competition is largely responsible for the pattern of environmental correlations in damage. Environmental correlations were just as common as genetic correlations in resistance, but the environmental correlations tended to be lower in magnitude, were more likely to be negative, and were more evenly spread out among the herbivore community than the genetic correlations. Damage levels by all 11 species were negatively correlated with damage by at least one other species. Thus the selective advantage a plant would receive from increased resistance any of these herbivores would be partially negated by increased damage by competing herbivores. As a result, competition has the potential to be an important constraint on the evolution of resistance in S. carolinense.     

Age-biased parasitism and density-dependent distribution of fleas (Siphonaptera) on a desert rodent

Parasites often confront conflicting demands when evaluating and distributing themselves among host individuals, in order to attain maximum reproductive success. We tested two alternative hypotheses about host preference by fleas in relation to the age of their rodent host. The first hypothesis suggests that fleas select adult over juvenile rodents because the latter represent a better nutritional resource (the “well-fed host” hypothesis), whereas the second hypothesis suggests that fleas prefer the weaker and less resistant juveniles because they are easier to colonise and exploit (“poorly fed host” hypothesis). We sampled fleas (Synosternus cleopatrae) on the gerbil (Gerbillus andersoni) in 23 different plots in the Negev desert and found an unequal distribution of fleas between adult and juvenile hosts. Furthermore, flea distribution changed as a function of flea density—from juvenile-biased flea parasitism (the “poorly fed host” hypothesis) at low densities to adult-biased flea parasitism (the “well-fed host” hypothesis) at high densities. Other factors that influenced flea preference were soil temperature and the presence of ticks. These results suggest that host selection is not an explicit alternative choice between adults and juveniles (“well-fed host” versus “poorly fed host” hypotheses), but rather a continuum where the distribution between adults and juveniles depends on host, parasite, and environmentally related factors.     

The effects on terrestrial arthropod communities of invasion of a coastal heath ecosystem by the exotic weed bitou bush (<Emphasis Type="Italic">Chrysanthemoides monilifera</Emphasis> ssp. <Emphasis Type="Italic">rotundata</Emphasis> L.)

Bitou bush (Chrysanthemoides monilifera ssp. rotundata) is a major environmental weed of littoral habitats on the southeastern coast of Australia. This study investigates the impacts upon selected arthropod assemblages of habitat invasion by this weed. Sixteen sites were placed at four geographic localities within nature reserves between Forster and Budgewoi on the coast of New South Wales. The sampling design included two spatial scales (between and within localities) and eight repeat samples (taken at two scales of temporal separation). Arthropods were collected from both arboreal and epigaeic micro-habitats. Specimens of ants (Hymenoptera: Formicidae), beetles (Coleoptera), true bugs (Heteroptera) and spiders (Araneae) were identified to species level. Differences in α diversity and species abundance distributions between the taxonomic assemblages are described, along with comparisons of data contrasts between bitou bush-free (“control”) and bitou bush-invaded (“impact”) habitats and between geographic localities. A subsidiary impact associated with herbicide application for control of the weed is also examined. Analysis of environmental differences indicates that bitou bush acts as a dampening agent, reducing the degree of seasonal fluctuation in factors such as leaf litter cover. Arboreal Heteroptera were the only group to show consistent evidence of significant differences in taxon richness or abundance between control and impact treatments within a locality, seen as a disruption of normal seasonal variation in diversity in bitou bush-impacted sites. Significant differences between geographic localities were more common, suggesting an effect at broader spatial scales. Evidence for arthropod assemblages characteristic of specific vegetation types was detected for several groups, as were changes in arthropod assemblage composition following application of herbicide for bitou bush control.     

Impact of two specialist insect herbivores on reproduction of horse nettle,Solanum carolinense

The frequency of coevolution as a process of strong mutual interaction between a single plant and herbivore species has been questioned in light of more commonly observed, complex relationships between a plant and a suite of herbivore species. Despite recognition of the possibility of diffuse coevolution, relatively few studies have examined ecological responses of plants to herbivores in complex associations. We studied the impact of two specialist herbivores, the horse nettle beetle, Leptinotarsa juncta, and the eggplant flea beetle, Epitrix fuscula, on reproduction of their host, Solanum carolinense. Our study involved field and controlled-environment experimental tests of the impact on sexual and potential asexual reproduction of attack by individuals of the two herbivore species, individually and in combination. Field tests demonstrated that under normal levels of phytophagous insect attack, horse nettle plants experienced a reduction in fruit production of more than 75% compared with plants from which insects were excluded. In controlled-environment experiments using enclosure-exclosure cages, the horse nettle's two principal herbivores, the flea beetle and the horse nettle beetle, caused decreases in sexual reproduction similar to those observed in the field, and a reduction in potential asexual reproduction, represented by root biomass. Attack by each herbivore reduced the numbers of fruits produced, and root growth, when feeding in isolation. When both species were feeding together, fruit production, but not root growth, was lower than when either beetle species fed alone. Ecological interactions between horse nettle and its two primary herbivores necessary for diffuse coevolution to occur were evident from an overall analysis of the statistical interactions between the two herbivores for combined assessment of fruit and vegetative traits. For either of these traits alone, the interactions necessary to promote diffuse coevolution apparently were lacking.     

The notoriously destructive potato stalk borer (<Emphasis Type="Italic">Trichobaris trinotata</Emphasis>) has negligible impact on its native host, <Emphasis Type="Italic">Solanum carolinense</Emphasis> (horsenettle)

While herbivores can have a wide range of impacts on their host plants, instances in which herbivore damage is truly beneficial are rare. Thus, it was surprising that damage by the potato stalk borer (PSB) was associated with increased fitness of its native host plant (horsenettle) in a recent field study, particularly because this insect can have such severe negative effects on potato crops. A controlled experiment with potted horsenettle was conducted to investigate the impact of damage by PSB to see if it has any negative impacts on the plants, or whether horsenettle truly overcompensates for its damage. Damage by PSB had negligible impact on sexual reproduction and potential future vegetative propagation through root production in this experiment. Additionally, PSB damage did not exacerbate the negative fitness impact of heavy leaf damage by the eggplant lace bug. The lack of positive impacts in the potted horsenettle study suggests that selection for decreased resistance to PSB in the field study was due to correlated selection for increased plant vigor, rather than to overcompensation. That is, when given options, PSB females may preferentially oviposit on larger, healthier plants that are likely to be better hosts for their larvae. An important implication of these results is that a wild weed’s high tolerance of a natural herbivore can compound the weed’s potential to serve as a reservoir from which the herbivore can switch to closely related agricultural plants that are not tolerant of its damage.     

The effect of rhizosphere microbes outweighs host plant genetics in reducing insect herbivory

Rhizosphere microbes affect plant performance, including plant resistance against insect herbivores; yet, a direct comparison of the relative influence of rhizosphere microbes versus plant genetics on herbivory levels and on metabolites related to defence is lacking. In the crucifer Boechera stricta, we tested the effects of rhizosphere microbes and plant population on herbivore resistance, the primary metabolome, and select secondary metabolites. Plant populations differed significantly in the concentrations of six glucosinolates (GLS), secondary metabolites known to provide herbivore resistance in the Brassicaceae. The population with lower GLS levels experienced ~60% higher levels of aphid (Myzus persicae) attack; no association was observed between GLS and damage by a second herbivore, flea beetles (Phyllotreta cruciferae). Rhizosphere microbiome (disrupted vs. intact native microbiome) had no effect on plant GLS concentrations. However, aphid number and flea beetle damage were respectively about three‐ and seven‐fold higher among plants grown in the disrupted versus intact native microbiome treatment. These differences may be attributable to shifts in primary metabolic pathways previously implicated in host defence against herbivores, including increases in pentose and glucoronate interconversion among plants grown with an intact microbiome. Furthermore, native microbiomes with distinct community composition (as estimated from 16s rRNA amplicon sequencing) differed two‐fold in their effect on host plant susceptibility to aphids. The findings suggest that rhizosphere microbes, including distinct native microbiomes, can play a greater role than population in defence against insect herbivores, and act through metabolic mechanisms independent of population.     

Development of SCAR markers linked to powdery mildew (<Emphasis Type="Italic">Uncinula necator</Emphasis>) resistance in grapevine (<Emphasis Type="Italic">Vitis vinifera</Emphasis> L. and <Emphasis Type="Italic">Vitis</Emphasis> sp.)

Sequence-characterized amplified regions markers (SCARs) were developed from six randomly amplified polymorphic DNA (RAPD) markers linked to the major QTL region for powdery mildew (Uncinula necator) resistance in a test population derived from the cross of grapevine cultivars “Regent” (resistant) × “Lemberger”(susceptible). RAPD products were cloned and sequenced. Primer pairs with at least 21 nucleotides primer length were designed. All pairs were tested in the F1 progeny of “Regent” × “Lemberger”. The SCAR primers resulted in the amplification of specific bands of expected sizes and were tested in additional genetic resources of resistant and susceptible germplasm. All SCAR primer pairs resulted in the amplification of specific fragments. Two of the SCAR markers named ScORA7-760 and ScORN3-R produced amplification products predominantly in resistant individuals and were found to correlate to disease resistance. ScORA7-760, in particular, is suitable for marker-assisted selection for powdery mildew resistance and to facilitate pyramiding powdery mildew resistance genes from various sources.     

The evolution of plant response to herbivory: simultaneously considering resistance and tolerance in Brassica rapa

Although the evolution of plant response to herbivory can involve either resistance (a decrease in susceptibility to herbivore damage) or tolerance (a decrease in the per unit effect of herbivory on plant fitness), until recently few studies have explicitly incorporated both of these characters. Moreover, theory suggests these characters do not evolve independently, and also that the pattern of natural selection acting on resistance and tolerance depends on their costs and benefits. In a genotypic selection analysis on an experimental population of Brassica rapa (Brassicaceae) I found a complex set of correlational selection gradients acting on resistance and tolerance of damage by flea beetles (Phyllotreta cruciferae: Chrysomelidae) and weevils (Ceutorhynchus assimilis: Curculionidae), as well as directional and stabilizing selection on resistance to attack by weevils. Evolution of response to flea beetle attack is constrained by a strong allocation cost of tolerance, and this allocation cost may be caused by a complex correlation among weevil resistance, weevil tolerance, flea beetle resistance, and flea beetle tolerance. Thus, one important conclusion of this study is that ecological costs may involve complex correlations among multiple characters, and for this reason these costs may not be detectable by simple pairwise correlations between characters. The evolution of response to weevil attack is probably constrained by a series of correlations between weevil resistance, weevil tolerance, and fitness in the absence of weevil damage, and possibly by a cost of tolerance of weevil damage. However, the nature of these constraints is complicated by apparent overcompensation for weevil damage. Because damage by both flea beetles and weevils had non-linear effects on plant fitness, standard measures of tolerance were not appropriate. Thus, a second important contribution of this study is the use of the area under the curve defined by the regression of fitness on damage and damage-squared as a measure of tolerance. This revised version was published online in July 2006 with corrections to the Cover Date.     

Dung beetle distribution patterns in the Iberian peninsula

A canonical variate analysis was used to summarize the distribution pattern of 30 species of dung beetles (Scarabaeinae) from 162 sites in the Iberian peninsula (Spain and Portugal) in relation to 4 site variables. The best separation of beetle species, obtained using a xerothermic climatic index, was into 3 groups called “mediterranean”, “unrestricted” and “temperate”. Little further separation was achieved using altitude, dung type or soil type. These data are used to select dung beetles for introduction to south-western Australia for bush fly control.

---

Résumé On a fait appel à l'analyse de variables canoniques pour résumer le plan de distribution, relatif à 4 variables de localité, de 30 espèces de bousiers (Scarabaeinae) de 162 localités de la Péninsule ibérique (l'Espagne et le Portugal). La meilleure division des espèces, obtenue à l'aide d'un indice xérothermique climatique, se faisait en 3 groupes désignés respectivement comme “méditerranéen”, “ubiquiste” et “tempéré”. Peu de séparation supplémentaire a été obtenue en ayant recours à des facteurs d'altitude, de nature de la bouse ou de type de sol. Ces données ont été employées pour aider à la sélection de bousiers destinés à être introduits en Australie Occidentale pour y combattreMusca vetustissima Walker[Diptera].

     

Impacts of simulated livestock grazing on Utah prairie dogs (Cynomys parvidens) in a low productivity ecosystem

Allometric foraging theory suggests that herbivores of greatly differing size should co-exist through niche segregation, but a few studies of large–small herbivore foraging relationships have reported competitive interactions. This study addresses the potential roles of habitat productivity and large herbivore grazing intensities on large–small herbivore foraging interactions. We examined effects of different intensity simulated grazing treatments on forage abundance and quality for Utah prairie dogs (Cynomys parvidens) in a low productivity ecosystem, and consequent effects on prairie dog individual growth rates, foraging preferences, and activity budgets. We hypothesized that simulated grazing would have predominantly facilitative impacts on Utah prairie dogs, as was found for black-tailed prairie dogs in higher productivity ecosystems. To test this hypothesis, we measured the effects of simulated grazing on forage nitrogen, digestibility, and biomass. Simulated grazing increased average forage nitrogen and digestibility while decreasing forage biomass. These effects were associated with reduced individual growth rates, increased juvenile foraging time, and reduced juvenile vigilance. Results suggest that the negative effects of reduced vegetation biomass greatly outweighed positive treatment effects in this study. However, prairie dogs in the moderate intensity defoliation treatment showed some preference for “grazed” plots over “ungrazed” plots, and this preference increased with time. Our study lends support to the idea that habitat productivity and herbivore densities may mediate shifts between facilitative and competitive interactions between different-sized herbivores.     

The scale of landscape fragmentation affects herbivore response to vegetation heterogeneity

Using alternating bands of weeds and broccoli I experimentally manipulated vegetation composition and the spatial scale at which the landscape was fragmented in a factorial design. This experimental approach allowed me to distinguish the effect of spatial scale from that of simple crop heterogeneity on crop herbivores. The importance of scale depended on which insect species were examined. Cabbage aphids (Brevicoryne brassicae) were influenced by vegetation composition at all tested scales of fragmentation; cabbage butterflies (Pieris rapae) were not affected by scale or by composition and flea beetles (Phyllotreta cruciferae) revealed a striking dependence on scale of fragmentation as well as an interaction between scale and composition. This approach shows the importance of dissecting out the effects of scale from other aspects of landscape manipulation, and emphasizes the challenge of developing a theory that will enable prediction of species–specific responses to scale. Received: 15 February 1998 / Accepted: 10 June 1998     

Inheritance of resistance to Plum pox virus in apricot (Prunus armeniaca L.)

The inheritance of resistance to Plum pox virus (PPV) has been studied in 1,178 apricot hybrids. Seven hundred and eighteen F1 hybrids, obtained from controlled crosses between the susceptible Greek cultivar “Bebecou” and the resistant PPV cultivars of American origin (“Stark Early Orange,” ‘NJA2,” ‘Veecot,” “Sunglo,” “Harlayne,” and “Orangered”) were evaluated for resistance to the PPV-M (Marcus) strain, 8 years after artificial inoculation. The inheritance of resistance to PPV has been additionally studied for the first time in a BC₁ population of 95 apricot hybrids for four vegetative periods. Reaction of each hybrid to PPV-M was scored through visual symptoms, indexing onto GF-305 and double-antibody sandwich enzyme-linked immunosorbent assay tests. Segregation within the hybrids, determined by Chi-squared analysis, fits a 1:1 ratio (P ≤ 0.05) of the resistant vs susceptible, indicating that resistance to PPV is controlled by a single dominant gene locus and that the above six resistant cultivars are heterozygous for the trait. Plants carrying this gene may initially develop disease symptoms on leaves but eventually recover and no virus can be detected in leaves. Susceptible plants show similar symptoms initially but remain symptomatic. Inheritance of resistance to PPV also has been studied in 365 F1 hybrids by crossing the resistant cultivar “Stella” with the susceptible “Bebecou” and the resistant cultivars “Sunglo” and “NJA2,” for 8 years after inoculation. The segregation ratio was 1:0 (resistant/susceptible) suggesting that “Stella” is homozygous for the resistance trait. The purpose of this work was the enhancement of the knowledge of inheritance of resistance to PPV for the selection of new cultivars.     

The evolution of “egalitarian” and “despotic” social systems among macaques

Recent studies of captive macaques have revealed considerable inter-species differences in dominance styles among females. In “egalitarian” species such as stumptail (Macaca arctoides) or tonkean macaques (M. tonkeana), social interactions are more symmetrical and less kin-biased than in “despotic” species such as Japanese (M. fuscata) or rhesus macaques (M. mulatta). Field observations of moor macaques (M. maurus), close relatives of tonkean macaques, suggest that tolerance during feeding characterizes their egalitarian dominance style in the natural habitat. Although it has been proposed that communal defense against other groups may be the main selective force in the evolution of egalitarian dominance style among females, few field data support this prediction. A game theory analysis showed that both an “egalitarian” strategy and a “despotic” strategy are possible evolutionarily stable strategies (ESS) under certain conditions. The difference in dominance styles might reflect the difference in ESS. This means that an egalitarian dominance style can emerge without strong between-group contest competition. A phylogenetic comparison among macaques suggests that despotic dominance styles very likely evolved from egalitarian dominance styles. In the future, primate socioecological studies should pay more attention to the evolutionary history of each species.     

Evolutionary ecology of resistance to herbivory: an investigation of potential genetic constraints in the multiple-herbivore community of Solanum carolinense

As part of a study of the ecological and evolutionary dynamics of herbivore resistance in Solanum carolinense (horsenettle), potential genetic constraints to the evolution of resistance to 11 of its most common herbivores were investigated. Leaf, flower, fruit, and stem herbivory were measured in a field experiment involving 24 ramets of each of 40 horsenettle genets. The experimental plant population contained significant genetic variation for resistance to all 11 species of herbivore. For only one species was there an indication of a genotype-by-environment interaction in the expression of resistance that might constrain its evolution. Genetic correlations in resistance to different species were common but not universal, with seven negative and 12 positive correlations out of the 55 pairwise species comparisons. Correlations were independent of plant part fed upon. The evolution of the resistance of horsenettle to most of its diverse community of herbivores does not appear to be prevented by a lack of genetic variation or by genotype-by-environment interactions in resistance. Negative genetic correlations in resistance to different herbivores may play a small role in slowing the evolution of resistance, but positive correlations may play at least as large a role in facilitating its evolution.     

Polymorphism of canonical and noncanonical <Emphasis Type="Italic">gypsy</Emphasis> sequences in different species of <Emphasis Type="Italic">Drosophila </Emphasis><Emphasis Type="Italic">melanogaster</Emphasis> subgroup: possible evolutionary relations

Mobile genetic elements constitute a substantial part of eukaryotic genome and play an important role in its organization and functioning. Co-evolution of retrotransposons and their hosts resulted in the establishment of control systems employing mechanisms of RNA interference that seem to be impossible to evade. However, “active” copies of endogenous retrovirus gypsy escape cellular control in some cases, while its evolutionary elder “inactive” variants do not. To clarify the evolutionary relationship between “active” and “inactive” gypsy we combined two approaches: the analysis of gypsy sequences, isolated from G32 Drosophila melanogaster strain and from different Drosophila species of the melanogaster subgroup, as well as the study of databases, available on the Internet. No signs of “intermediate” (between “active” and “inactive”) gypsy form were found in GenBank, and four full-size G32 gypsy copies demonstrated a convergence that presumably involves gene conversion. No “active” gypsy were revealed among PCR generated gypsy ORF3 sequences from the various Drosophila species indicating that “active” gypsy appeared in some population of D. melanogaster and then started to spread out. Analysis of sequences flanking gypsy variants in G32 revealed their predominantly heterochromatic location. Discrepancy between the structure of actual gypsy sites in G32 and corresponding sequences in database might indicate significant inter-strain heterochromatin diversity. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.