Interspecific hybridization of plants and resistance to herbivores: hypotheses,genetics, and variable responses in a diverse herbivore community

We studied the morphology, molecular genetics, and hebivory of two species of willows (Salix sericea and S. eriocephala) and their interspecific hybrids to test four alternative hypotheses concerning the effects of hybridization on plant resistance. Individually marked plants were identified using morphological traits in the field and measurements of stipule and leaf pubescence were made and compared using Canonical Discriminant Function Analysis. DNA was extracted from the leaves of a sample of the marked plants and RAPD-PCR analysis was performed to establish the genetic status of parental and hybrid plants. RAPD band analysis generally verified the genetic status of parental plants. Hybrid plants were usually correctly identified in the field with a few exceptions. However, the hybrid plants were a heterogeneous group of plants made up of most plants that appear to be F₁s and a few plants that appear to be backcrosses to S. sericea. Morphological variables were useful for distinguishing S. sericea from S. eriocephala and hybrids, but were not as dependable in distinguishing between S. eriocephala and hybrids. We compared the densities of 11 herbivore species and the infection by a leaf rust pathogen (Melampsora sp.) on the leaves and stems of two parents and the hybrids in the field. We found support for the Additive hypothesis (3 species), the Dominance hypothesis (2 species) and the Hybrid Susceptibility hypothesis (7 species, 6 herbivores and the Melampsora rust). We found no evidence for the Hybrid Resistance hypothesis. Guild membership was not a good predictor of similar responses of species to hybrid versus parental plants. A Canonical Discriminant Function Analysis showed discrete separation of the taxa based on herbivore densities, illustrating different community structures on hybrid and parental plants. This study demonstrates the diversity of responses of phytophages in response to interspecific hybridization.     

Interspecific and temporal variation in herbivore responses to hybrid willows

We studied herbivory of two species of willows (Salix sericea and S. eriocephala) and their interspecific hybrids to test alternative hypotheses concerning the effects of hybridization on plant resistance. Individually marked plants were identified using morphological traits in the field and random amplified polymorphic DNA (RAPD) band analysis was used to verify the genetic status of many parental and hybrid plants. The desities of 12 herbivore species on plants in the field were compared between two parents and their F₂-type hybrids. We found about equal support for the additive, dominance, and hybrid susceptibility hypotheses over 4 years. In one year, one species supported the hybrid resistance hypothesis. Guild membership was not a good predictor of similar responses of species to hybrid versus parental plants. There were marked differences in support for particular hypotheses among years for four herbivore species. This study demonstrates the diversity of responses of phytophages in response to interspecific hybridization, and indicates that year-to-year variation in relative resistance of hybrid plants can be important.     

Willow hybrids and herbivory: a test of hypotheses of phytophage response to hybrid plants using the generalist leaf-feeder Lochmaea caprea (Chrysomelidae)

To test hypotheses of herbivore response to hybrid plants, I determined the preference of the generalist beetle Lochmeae capreae for individuals of Salix caprea, Salix phylicifolia and F1 hybrids between these species. F1 hybrids and pure individuals were created by hand-pollination of willow individuals in the field. The preference tests were performed in 1994 and 1995 and in both years there was a tendency for higher leaf area consumption from hybrids than from the parental species. However, statistically significant differences in consumption were only found in 1995 between hybrids and S. phylicifolia. The result from this study therefore lends most support to the dominance hypothesis: hybrid susceptibility was similar to that of one of the parental species. This is one of the first strict preference tests showing that a generalist herbivore can detect differences in palatability between hybrids and parental species. Furthermore, the experimental design used in the study minimized environmental variation. The results should therefore primarily reflect genetically based quality differences between hybrids and parental species. Received: 4 March 1996 / Accepted: 12 August 1996     

Genetic architecture of susceptibility to herbivores in hybrid willows

We performed a common garden experiment using parental, F1, F2, and backcross willow hybrids to test the hypothesis that hybrid willows experience breakdown of resistance to herbivores. After exposing plants to herbivores in the field, we measured the densities/damage caused by 13 insect herbivores and one herbivorous mite. Using joint-scaling tests, we determined the contribution of additive, dominance, and epistasis to variation in susceptibility to herbivores (measured either as density or damage level) among the six genetic classes. We found the genetic architecture of susceptibility/resistance in the parental species to be complex, involving additive, dominance, and epistasis for each herbivore species. Although genic interactions altered plant susceptibility for each of the 14 herbivores, three distinct patterns of response of herbivores to hybrids were expressed. One pattern, observed in four herbivore species, supported the hypothesis of breakdown of resistance genes in recombinant hybrids. A second pattern, shown by six other herbivore species, supported the hypothesis of hybrid breakdown of host recognition genes. In other words, epistatic interactions for host recognition traits (probably oviposition/feeding stimulants or attractants) appeared to be important in determining herbivore abundance for those six species. The final patterns supported a structure of dominance, either for host recognition traits (in the case of three herbivore species) or for host resistance traits (for one herbivore species). The combination of differing responses of herbivore species, including members of the same genus and tribe, and the ubiquitous importance of epistasis suggests that many genes affect herbivore resistance in this hybrid willow system.     

Plant genetic differences influence herbivore community structure: evidence from a hybrid willow system

To determine the influence of plant genetic variation on community structure of insect herbivores, we examined the abundances of 14 herbivore species among six genetic classes of willow: Salix eriocephala, S. sericea, their F₁ and F₂ interspecific hybrids, and backcross hybrids to each parental species. We placed 1-year-old plants, grown from seeds generated from controlled crosses, in a common garden. During the growing season, we censused gall-inducing flies and sawflies, leaf-mining insects, and leaf-folding Lepidoptera to determine the community structure of herbivorous insects on the six genetic classes. Our results provided convincing evidence that the community structure of insect herbivores in this hybrid willow system was shaped by genetic differences among the parental species and the hybrid genetic classes. Using MANOVA, we detected significant differences among genetic classes for both absolute and relative abundance of herbivores. Using canonical discriminant analysis, we found that centroid locations describing community structure of the insect herbivores differed for each genetic class. Moreover, the centroids for the four hybrid classes were located well outside of the range between the centroids for the parental species, suggesting that more than additive genetic effects of the two parental species influenced community formation on hybrid classes. Line-cross analysis suggested that plant genetic factors responsible for structuring the herbivore community involved epistatic effects, as well as additive and dominance effects. We discuss the ramifications of these results in regard to the structure of insect herbivore communities on plants and the implications of our findings for the evolution of interspecific interactions.     

Three-trophic-level interactions in cattail hybrid zones

Plant-herbivore and herbivore-parasitoid wasp interactions were examined in three hybrid zones of the cattails Typha latifolia and T. angustifolia in south-eastern Michigan over a 2-year period. Patterns of resource use by two lepidopteran species of seed-eating herbivores were studied and herbivore densities sustained by the hybrid cattail T. x glauca and its parental species were determined. Densities of the commoner seed-eating lepidopteran, Lymnaecia phragmitella, were found to be highest in seed heads of both parental species and lowest in hybrid seed heads in zones of hybridization, thus supporting the hybrid resistance hypothesis of Fritz et al. (1994). Densities of the second herbivore, Dicymolomia julianalis, on the hybrid were lower than on the parental T. latifolia, but did not differ from the mean of the combined herbivore densities of the two parental species. D. julianalis in cattails appears to fit the additive hypothesis (Fritz et al. 1994). Parasitism of L. phragmitella by four species of parasitoid wasp, Itoplectis conquisitor (Ichneumonidae), Scambus hispae (Ichneumonidae), Macrocentrus delicatus (Braconidae), and Temelucha gracilipes (Ichneumonidae), was studied to determine if rates of parasitization by these natural enemics on the different cattails were similar to the patterns of plant exploitation exhibited by their hosts. Parasitism rates of L. phragmitella were significantly higher in larvae reared on hybrid cattails. This study reveals two different patterns of hybrid utilization by two very similar herbivores. Opposite patterns of plant utilization were found for the parasitoids compared with their host (L. phragmitella) in cattail hybrid zones.     

Herbivore resistance of invasive <Emphasis Type="Italic">Fallopia</Emphasis> species and their hybrids

Hybridization has been proposed as a mechanism by which exotic plants can increase their invasiveness. By generating novel recombinants, hybridization may result in phenotypes that are better adapted to the new environment than their parental species. We experimentally assessed the resistance of five exotic Fallopia taxa, F. japonica var. japonica, F. sachalinensis and F. baldschuanica, the two hybrids F. × bohemica and F. × conollyana, and the common European plants Rumex obtusifolius and Taraxacum officinale to four native European herbivores, the slug Arion lusitanicus, the moth Noctua pronuba, the grasshopper Metrioptera roeselii and the beetle Gastrophysa viridula. Leaf area consumed and relative growth rate of the herbivores differed significantly between the Fallopia taxa and the native species, as well as among the Fallopia taxa, and was partly influenced by interspecific variation in leaf morphology and physiology. Fallopia japonica, the most abundant Fallopia taxon in Europe, showed the highest level of resistance against all herbivores tested. The level of resistance of the hybrids compared to that of their parental species varied depending on hybrid taxon and herbivore species. Genotypes of the hybrid F. × bohemica varied significantly in herbivore resistance, but no evidence was found that hybridization has generated novel recombinants that are inherently better defended against resident herbivores than their parental species, thereby increasing the hybrid’s invasion success. In general, exotic Fallopia taxa showed higher levels of herbivore resistance than the two native plant species, suggesting that both parental and hybrid Fallopia taxa largely escape from herbivory in Europe.     

Seasonal dynamics of dry weight, growth rate and root/shoot ratio in different aged seedlings of Salix caprea

Willows (e.g. Salix caprea L.) are deciduous and richly branched shrubs or small trees. Salix caprea shows a high adaptability to different habitat conditions. One way of evaluating this adaptability is to measure willow biomass and production. Young plants of S. caprea were sampled from the bottom of an artificial lagoon in which sediments removed from the local Vajgar fishpond were deposited. The bottom of the lagoon was overgrown by vegetation dominated by seedlings of the willows S. caprea and S. aurita. Willows grew in the lagoon at average density of 58 plants per m². The biomass production and growth of S. caprea were determined for 15 samples (collected from 315 individuals) during the growing season. Annual net dry matter production in the whole community was estimated for 2.7 kg m⁻². Willows are generally considered to be fast-growing plants. The highest RGR of willows recorded by us was 0.03 to 0.04 g g⁻¹ d⁻¹ both in the stems and roots. This value was often recorded from July to August.     

Alternative seed defence mechanisms in a palo verde (Fabaceae) hybrid zone: effects on bruchid beetle abundance

Abstract.

• 1 We asked three questions about the patterns of relative abundance of insect herbivores across host plant taxa at a palo verde hybrid zone. (1) What is the morphological structure of the hybrid zone and does this suggest a certain pattern of introgression? (2) How are putative parental seed defence mechanisms expressed in hybrid plants? (3) Do ovipositing females prefer host plant taxa on which their offspring have best survivorship?
• 2 Morphologically, hybrids were either intermediate or tended to resemble one parental species. Previous studies have suggested that unidirectional introgression results in loss of parental defence mechanisms against herbivores. Hybrid plants in general lacked seed coat resistance and early pod abscission which are known to act as plant defence mechanisms against bruchid beetles in the parental palo verde trees.
• 3 All other sources of bruchid mortality that we examined did not vary across parental and hybrid taxa, with the possible exception of egg parasitism which occurred at a lower frequency on one parental palo verde species.
• 4 Thus, survivorship of bruchid offspring should be greater on hybrid palo verdes.
• 5 Patterns of egg densities suggested that females may prefer hybrid hosts in some years but not others. An oviposition choice experiment conducted in the field, however, showed bruchids have no preference for hybrids over one of the parental species.
• 6 These results suggest that some insect herbivores may have higher densities on hybrid host plants because they are less resistant.

     

Tritrophic interactions and trade-offs in herbivore fecundity on hybridising host plants

Abstract. 1. Interspecific plant hybridisation can have important evolutionary consequences for hybridising plants and for the organisms that they interact with on multiple trophic levels. In this study the effects of plant hybridisation on the abundance of herbivores and on the levels of herbivore parasitism were investigated. 2. Borrichia frutescens, B. arborescens, and their hybrid (B. × cubana) were censused for Asphondylia borrichiae galls and Pissonotus quadripustulatus plant hoppers in the Florida Keys. Levels of egg parasitism were determined by dissecting parental and hybrid stems and galls for herbivore and parasite eggs and larvae. Stem toughness and gall size are plant‐mediated modes of protection from parasitism and these were also measured. For gall midges, fly size was measured as an estimate of fecundity. 3. Field censuses indicated that herbivore abundances varied on hybrid hosts relative to parent plant species and that the different herbivore species exhibited different patterns of abundance. Asphondylia borrichiae gall numbers followed the additive pattern of abundance while P. quadripustulatus numbers most closely resembled the dominance pattern. 4. Parasitism of P. quadripustulatus eggs was high on B. frutescens and the hybrids, and low on B. arborescens, which also had significantly tougher stems. Asphondylia borrichiae suffered the highest levels of parasitism on B. frutescens, the host plant which produced the smallest galls. On B. arborescens, which produced the largest galls, levels of A. borrichiae parasitism were lowest. Both parasitism and gall size were intermediate on the hybrid plants. Galls on B. arborescens and hybrid plants produced significantly smaller flies then those from B. frutescens suggesting that, when selecting hosts from among parent species and hybrids, gall flies may face a trade‐off between escape from natural enemies and maximising fecundity.     

Rust disease (Melampsora epited) of willow (Salix spp.) grown as short rotation coppice (SRC) in inter- and intra-species mixtures

Twenty inter‐ and intra‐species genotypes of willows (Salix spp.) were grown in large mono‐plots and incorporated into five, 10,15 and 20–way mixtures. In each growing season from 1996 to 2001 the level of rust disease caused by Melampsora epitea was recorded on each genotype where it was growing as a mono‐plot or as part of a mixture. Three genotypes, S. schwerinii×viminalis×dasyclados’V7531′, S. schwerinii×aquatica‘V7533’ and S. viminalis‘Gigantea’ remained virtually rust free through the six yr of the trial. It is, however, argued that there are benefits of including such genotypes in a mixture in order to reduce the risk of them developing rust susceptibility. S. burjatica‘Germany’, S. dasyclados×aquatica‘V7511’ and S. dasyclados×caprea‘V794’ were severely affected by rust, although levels tended to be less when included in mixtures. The levels of rust on S. mollissima‐undulata‘SQ83’ were much higher in the first three‐year harvest cycle than during the second cycle. On a number of genotypes, e.g. S. burjatica‘Germany’ and S. dasyclados×aquatica‘V7511’ rust was more severe on the regrowth from freshly coppiced stools.     

Drought changes plant chemistry and causes contrasting responses in lepidopteran herbivores

Drought events are predicted to increase due to climate change, yet consequences for plant–insect interactions are only partially understood. Drought‐mediated interactions between herbivores and their host plants are affected by a combination of factors, including characteristics of the affected plant, its associated herbivore and of the prevailing drought. Studying the effect of these factors in combination may provide important insight into plant and herbivore responses to drought. We studied drought effects on plant resistance to two leaf‐chewing herbivores by considering differing growth conditions, plant chemistry and insect responses in concert. We exposed Alliaria petiolata plants from several wild populations to different intensities of intermittent drought stress and quantified drought‐mediated changes in plant chemistry. Simultaneously, we assessed behavior (feeding preference) and performance of two lepidopteran herbivores: Pieris brassicae, a specialist, and Spodoptera littoralis, a generalist. Drought led to lowest concentrations of secondary defense compounds in severely stressed plants, without affecting total nitrogen content. Additionally, drought evoked opposite patterns in feeding preferences (plant palatability) between the herbivore species. Pieris brassicae consumed most of well‐watered plants, while S. littoralis preferred severely drought‐stressed plants. Hence, feeding preferences of S. littoralis reflected changes in plant secondary chemistry. Contrary to their feeding preference, P. brassicae performed better on drought‐stressed than on well‐watered plants, with faster development and higher attained pupal mass (plant suitability). Spodoptera littoralis showed retarded development in all treatments. In conclusion, drought caused plant secondary defense compounds to decrease consistently across all studied plant populations, which evoked contrasting feeding preferences of two herbivore species of the same feeding guild. These results suggest herbivore specificity as a possible explanation for herbivore responses to drought and emphasize the importance of herbivore characteristics such as feeding specialization in understanding and predicting consequences of future drought events.     

Interactions between willows and insect herbivores under enhanced ultraviolet-B radiation

We studied the effects of elevated ultraviolet-B radiation on interactions between insect herbivores and their host plants by exposing two species of phytochemically different willows, Salix myrsinifolia and S. phylicifolia, to a modulated increase in ultraviolet radiation in an outdoor experiment and monitoring the colonisation of insect herbivores on these willows. We examined the effect of increased ultraviolet-B (UV-B) radiation on (1) the quality of willow leaves, (2) the distribution and abundance of insect herbivores feeding on these willows, (3) the resulting amount of damage, and (4) the performance of insect larvae feeding on the exposed plant tissue. Six clones of each of the two willow species were grown in eight blocks for 12 weeks in the UV-B irradiation field. The clones were exposed to a constant 50% increase in UV-B radiation (simulating 20-25% ozone depletion), to a small increase in UV-A radiation or to ambient solar irradiation. We allowed colonisation on the willows by naturally occurring insects, but also introduced adults of a leaf beetle, Phratora vitellinae, a specialist herbivore on S. myrsinifolia. Increased UV-B radiation did not affect any of the measured indices of plant quality. However, numbers of P. vitellinae on S. myrsinifolia were higher in plants with UV-B treatment compared with UV-A and shade controls. In laboratory tests, growth of the second-instar larva of P. vitellinae was not affected by UV-B treatment of S. myrsinifolia, but was retarded on UV-B treated leaves of S. phylicifolia. In addition, naturally occurring insect herbivores were more abundant on willows exposed to elevated UV-B radiation compared to those grown under control treatments. In spite of the increased abundance of insect herbivores, willows treated with elevated UV-B did not suffer more herbivore damage than willows exposed to ambient solar radiation (shade control). The observed effects of UV-B on herbivore abundance, feeding and growth varied significantly due to spatial variation in environment quality, as indicated by the UV-treatment x block interaction. The results suggest that (1) environmental variation modifies the effects of UV-B radiation on plant-insect interactions and (2) specialist herbivores might be more sensitive to chemical changes in their secondary host plants (S. phylicifolia) than to changes in their primary hosts (S. myrsinifolia).     

Susceptibility and tolerance in hybrid and pure‐line rice varieties to herbivore attack: biomass partitioning and resource‐based compensation in response to damage

Hybrid rice has been noted for its susceptibility to insects and diseases compared to pure‐line (conventional) rice varieties. We investigated herbivory by Nilaparvata lugens, Sogatella furcifera and Scirpophaga incertulas on replicated three‐line hybrid sets (parental and hybrid lines) in field and greenhouse experiments. In a field experiment, caterpillar densities and stemborer damage was similar among hybrid and parental lines. In field and greenhouse experiments, the cytoplasmic male sterile (CMS)‐lines and maintainer lines had higher densities of planthoppers (including N. lugens and S. furcifera) than restorer or hybrid lines likely because of their wild abortive CMS‐lineage. High nitrogen levels increased plant mortality due to N. lugens, but often reduced mortality from S. furcifera and S. incertulas: this was similar between hybrid and pure‐line varieties. The hybrids were generally more tolerant of herbivory (lower biomass reductions per unit weight of insect) than the inbred parental lines. The addition of nitrogen to both the hybrid and pure‐line varieties had contrasting effects on tolerance depending on the nature of the attacking insect: fertiliser increased tolerance to S. furcifera (lower losses of yield and shoot biomass per mg insect) and S. incertulas (lower yield, shoot and root biomass loss) but fertiliser reduced tolerance to N. lugens (higher loss of root biomass and no effects on yield and shoot biomass loss). Our results indicate that hybrid rice is not physiologically more susceptible to herbivores than are pure‐line varieties even under high nitrogen conditions, but does have higher tolerance to insect damage.     

Herbivores and pathogens on willow: do they affect each other?

Abstract 1 Willows often need to cope with attack by both rust fungi and herbivores. We studied whether rust infection on willow affects the herbivore, and vice versa, whether herbivore feeding affects the fungal infection. The system investigated by laboratory bioassays and greenhouse experiments consisted of the willow hybrid Salix × cuspidata, the rust Melampsora allii‐fragilis and the willow leaf beetle Plagiodera versicolora. Effects were studied both on a local scale (rust infection and feeding on the same leaf) and systemically (rust infection and feeding on different, but adjacent leaves). 2 Rust infection was not affected by herbivore feeding on a local scale. Systemically, however, the willow's susceptibility towards rust infection was increased by herbivore feeding, as indicated by a higher number of rust sori on leaves adjacent to feeding‐damaged leaves. The herbivore's performance was detrimentally affected by rust infection: increase of mortality (systemically), decrease of larval weight (locally and systemically) and prolonging of developmental time (locally and systemically). 3 Previous rust infection enhanced systemically the willow's susceptibility towards subsequent fungal infection. Previous herbivore feeding on the willow had no effects on the herbivore's developmental time and mortality. However, feeding upon previously feeding‐damaged willow leaves significantly reduced larval weight.     

Phenolic glycosides and condensed tannins in Salix sericea, S. eriocephala and their F1 hybrids: not all hybrids are created equal

The performance of hybrids depends upon the inheritance and expression of resistance traits. Secondary chemicals are one such resistance trait. In this study, we measured the concentrations of phenolic glycosides and condensed tannins in parental and F1 hybrid willows to examine the sources of chemical variation among hybrids. S. sericea produces phenolic glycosides, salicortin and 2'-cinnamoylsalicortin, and low concentrations of condensed tannin in its leaves. In contrast, S. eriocephala produces no phenolic glycosides but high concentrations of condensed tannins in its leaves. These traits are inherited quantitatively in hybrids. On average, F1 hybrids are intermediate for condensed tannins, suggesting predominantly additive inheritance or balanced ambidirectional dominance of this defensive chemical from the parental species. In contrast, the concentration of phenolic glycosides is lower than the parental midpoint, indicating directional dominance. However, there is extensive variation among F1 hybrids. The concentration of tannin and phenolic glycosides in F1 hybrid families is either (1) lower than the midpoint, (2) higher than the midpoint, or (3) indistinguishable from the midpoint of the two parental taxa. It appears that the production of the phenolic glycosides, especially 2'-cinnamoylsalicortin, is controlled by one or more recessive alleles. We also observed a two-fold or greater difference in concentration between some hybrid families. We discuss how chemical variation may effect the relative susceptibility of hybrid willows to herbivores.     

Evolution of a hybrid zone of two willow species (Salix L.) in the European Alps analyzed by RAD-seq and morphometrics

Natural hybridization of plants can result in many outcomes with several evolutionary consequences, such as hybrid speciation and introgression. Natural hybrid zones can arise in mountain systems as a result of fluctuating climate during the exchange of glacial and interglacial periods, where species retract and expand their territories, resulting in secondary contacts. Willows are a large genus of woody plants with an immense capability of interspecific crossing. In this study, the sympatric area of two diploid sister species, S. foetida and S. waldsteiniana in the eastern European Alps, was investigated to study the genomic structure of populations within and outside their contact zone and to analyze congruence of morphological phenotypes with genetic data. Eleven populations of the two species were sampled across the Alps and examined using phylogenetic network and population genetic structure analyses of RAD Seq data and morphometric analyses of leaves. The results showed that a homoploid hybrid zone between the two species was established within their sympatric area. Patterns of genetic admixture in homoploid hybrids indicated introgression with asymmetric backcrossing to not only one of the parental species but also one hybrid population forming a separate lineage. The lack of F1 hybrids indicated a long-term persistence of the hybrid populations. Insignificant isolation by distance suggests that gene flow can act over large geographical scales. Morphometric characteristics of hybrids supported the molecular data and clearly separated populations of the parental species, but showed intermediacy in the hybrid zone populations with a bias toward S. waldsteiniana. The homoploid hybrid zone might have been established via secondary contact hybridization, and its establishment was fostered by the low genetic divergence of parental species and a lack of strong intrinsic crossing barriers. Incomplete ecological separation and the ability of long-distance dispersal of willows could have contributed to the spatial expansion of the hybrid zone.     

Differential effects of host plant hybridization on herbivore community structure and grazing pressure on forest canopies

Interspecific hybridization in plants is known to have ecological effects on associated organisms. We examined the differences in insect herbivore community structure and grazing pressure on tree canopy leaves among natural hybrids and their parental oak species. We measured leaf traits, herbivore community structure, and grazing pressure on leaves of two oak species, Quercus crispula and Q. dentata, and their hybrids. The concentration of nitrogen in canopy leaves was greater in hybrids and in Q. dentata than in Q. crispula. The concentration of total phenolics was lower in hybrids than in Q. crispula. The concentration of condensed tannin was greater in hybrids than in Q. crispula. Relative herbivore abundance and species richness were greater on oak hybrids than on either parental species; herbivore species diversity and composition on hybrids were close to those on Q. crispula. Herbivore grazing pressure was lower on hybrids and Q. dentata than on Q. crispula. There was a negative correlation between herbivore grazing pressure and leaf nitrogen, suggesting that interspecific variation among oak taxa in herbivore pressure may be explained by leaf nitrogen; variation in herbivore community structure among oak taxa is likely to be controlled by polygenic leaf traits. Differing responses of (1) herbivore community structure and (2) herbivore grazing pressure to host plant hybridization may play important roles in regulating herbivore biodiversity in cool‐temperate forest canopies.     

桉树杂交种对桉树枝瘿姬小蜂的抗性变异分析

植物种间杂交是一种普遍自然现象,杂交往往造成植物表型及生理变异,从而改变杂种抗虫性。与亲本种相比,杂种抗虫性可能增强或减弱,也有可能处于与亲本相似水平。初生、次生代谢物的质变与量变是引起杂种抗虫性变异的重要原因。近年来,桉树杂交育种已在世界范围内广泛应用并取得了显著成效,桉树杂交种间抗虫性表现参差不齐,因此,桉树是研究杂交种抗虫性变异机制的理想材料。以2个桉树杂交种巨细桉DH201-2、巨尾桉G9及桉树重要害虫桉树枝瘿姬小蜂为研究对象,比较了2个杂交种与其纯亲本种[(巨桉×细叶桉),(巨桉×尾叶桉)]间的抗虫性差异;同时,综合比较了品系间叶片性状(叶片厚度、含水率、比叶面积)、初生化合物(C、N、可溶性糖、可溶性蛋白)及次生化合物(总酚、单宁)差异,以研究桉树杂交种抗虫性变异的理化机制。结果表明:DH201-2感染桉树枝瘿姬小蜂的虫瘿数目显著高于其双亲本种,而G9上虫瘿数目显著低于其双亲本种。DH201-2与G9的叶片厚度与巨桉相近,而显著薄于另一亲本种。DH201-2叶片含水率显著高于细叶桉、与巨桉相近;G9叶片含水率则显著低于其双亲本种。相似的是,DH201-2和G9的比叶面积均显著高于其双亲本种。初生化合物方面,DH201-2叶片可溶性糖和可溶性蛋白含量均显著高于其亲本种,N含量则仅高于细叶桉;而G9叶片可溶性蛋白含量虽高于其双亲本种,可溶性糖含量则无显著差异,N含量显著低于其双亲本种。次生化合物方面,DH201-2叶片总酚和单宁含量显著低于其双亲本种,而G9则显著高于其双亲本种。因此,与其亲本种相比,DH201-2感虫性增加,而G9抗虫性增加;与桉树枝瘿姬小蜂发育相关的营养指标(如含水率、可溶性糖、N含量)及次生防御物质(如总酚、缩合单宁)在桉树杂交种组织内的含量差异影响了桉树杂交种对桉树枝瘿姬小蜂的抗性。在全球推行桉树杂交育种且桉树害虫数量逐年增加的大背景下,应加强对桉树杂交种抗虫性机制研究,为选育高抗品系及桉树产业可持续发展提供理论指导。