Diffuse selection on resistance to deer herbivory in the ivyleaf morning glory, Ipomoea hederacea

Recent work defines coevolution between plants and herbivores as pairwise when the pattern of selection on resistance traits and the response to selection are both independent of the presence or absence of other herbivores. In addition, for a pairwise response to selection, resistance to a focal herbivore must have the same genetic basis in the presence and absence of other herbivores. None of these conditions were satisfied for the ivyleaf morning glory, Ipomoea hederacea, and its insect, fungal, and mammalian natural enemies with a quantitative genetics field experiment. A significant negative genetic correlation exists between resistance to deer and generalist insect herbivory that would preclude an independent response to selection. In addition, resistance loci under selection differ depending on the composition of the natural enemy community as indicated by genetic correlations between deer resistances in the presence and absence of other natural enemies that differ substantially from 1. Finally, selection on deer resistance depends on the presence or absence of insects; in the presence of insects, greater deer resistance is favored, but in the absence of insects, deer resistance is effectively neutral. These results indicate that the composition of the natural enemy community can alter both the pattern of selection and the likely response to selection of resistance traits.     

THE EVOLUTION OF RESISTANCE TO HERBIVORY IN IPOMOEA PURPUREA. II. NATURAL SELECTION BY INSECTS AND COSTS OF RESISTANCE

An important component of the process of coevolution between plants and their insect herbivores is the imposition of selection on plants by insects. Although such selection has been inferred from indirect evidence, little direct evidence for it exists. One goal of this study was to seek direct evidence by determining, for a single plant-herbivore system, whether insect herbivores impose selection on their host plants. A second goal was to determine whether costs are associated with genotypes that confer resistance to herbivores, as has been commonly postulated. The annual morning glory, Ipomoea purpurea, exhibits genetic variation in resistance to four different types of insects. For three of these types, most of the genetic variation is additive. Removal of insect herbivores increased the number of seeds produced by I. purpurea by 20% and eliminated additive genetic variation for seed number (fitness). This result implies that herbivores impose selection on some trait(s) of their host plants. Coupled with selection for decreased damage by corn earworms, as revealed by a negative additive genetic covariance between damage and fitness, this result suggests that insect herbivores impose selection on resistance to corn earworms in I. purpurea. Two types of cost of resistance to herbivores were sought in I. purpurea: 1) internal trade-offs in allocation of resources and 2) ecological trade-offs between resistances to different insects. No costs of either type were detected. This result suggests that cost-benefit arguments that attempt to predict the evolution of levels of resistance to herbivores are not applicable to I. purpurea.     

An Ecological Genetic Approach to the Study of Coevolution

Almost forty years ago, Ehrlich and Raven (1964) hypothesizedthat the great diversity of plants and the herbivores that feedon them arose from a process of coevolution. Plants do possessan amazing diversity of traits that are easily imagined as havingarisen from an antagonistic interaction between plants and herbivores.Two basic assumptions lie at the root of most theories of coevolutionbetween plants and their herbivores. First, herbivores are agentsof natural selection on plant resistance traits. Second, plantsincur a significant fitness cost for possessing these resistancetraits. An ecological genetic approach can provide rigorousevidence for these coevolutionary assumptions. In this paper,I present new experimental work on the subject of costs of resistanceand review and discuss my own previous work bearing directlyon these questions. Using both field experiments on naturalpopulations of the mouse-ear cress (Arabidopsis thaliana) andlaboratory experiments using genetically modified plants, Idemonstrate that herbivores are exerting selection on both achemical and physical resistance trait and that there are significantfitness costs to possessing these two traits. These resultsprovide direct confirmation that our current models of the evolutionof plant defenses are appropriate.     

Differential attack on diploid, tetraploid, and hexaploid Solidago altissima L. by five insect gallmakers

Genetic variation among plants can influence host choice and larval performance in insect herbivores. Ploidy (cytotype) variation is a particularly dramatic form of plant genetic variation, and where diploid and polyploid cytotypes of a species occur in sympatry, they may provide herbivores with choices that are distinguished by profound and genome-wide genetic differences. We tested for non-random attack by five gallmaking insect herbivores on diploid, tetraploid, and hexaploid cytotypes of the goldenrod Solidago altissima L., working in seven midwestern US populations where the ploidies co-occur on spatial scales relevant to insect host choice. For four of the five herbivores, attack was non-random with respect to ploidy at one or more sites. Ploidy effects on attack were complex: the ploidy subjected to highest attack varied both across herbivores within sites and (for most herbivores) across sites within herbivores. Ploidy effects on attack will alter rates of encounter between insect herbivores—either increasing or decreasing the likelihood of two herbivores sharing a host plant ramet, compared with the case with no effects of ploidy. Plant ploidy variation appears likely to have a major impact on insect community organization, and perhaps on plant–herbivore coevolution, but that impact is likely to be spatially heterogeneous. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

植物与草食动物之间的协同适应及进化

通常协同进化是指一个物种 (或种群 )的遗传结构由于回应于另一个物种 (或种群 )遗传结构的变化而发生的相应改变。广义的理解 ,协同进化是相互作用的物种之间的互惠进化。生物之间、特别是植物与草食动物之间的协同适应与进化 ,已经成为生物进化、生态、遗传等学科十分关注的问题 ,可能成为生物学中各学科研究的交汇点或结点。作者具体阐述了 :(1)生物之间协同进化的研究意义 ,包括对生物学与生态学的价值 ;(2 )生物之间协同进化研究的限制或困难 ,诸如时间、研究对象、进化等级尺度和研究方法的限制 ;(3)植物与草食动物之间协同进化的主要研究对象 (系统 ) ,即昆虫传粉系统、昆虫诱导植物反应系统、种子散布系统、以及大型草食动物采食与植物反应系统 ;(4 )植物与草食动物之间协同进化的主要研究内容 ,包括适应特征 (性状 )——物种的可塑性 ,以及适应机制——物种适应过程与策略两个方面 ;(5 )植物与草食动物之间协同进化研究的存在问题及研究方向     

GENETICS OF RESISTANCE OF SALIX SERICEA TO A DIVERSE COMMUNITY OF HERBIVORES

We measured resistance of Salix sericea, the silky willow, to a diverse assemblage of 12 herbivores. We investigated the potential for multispecies coevolution among these herbivores by measuring genetic correlations between pairs of herbivores interacting within the component community. After measuring herbivore attack on half-sib families of potted S. sericea during three years, we found significant narrow-sense heritabilities of resistance to Phyllonorycter salicifoliella and Phyllocnistis sp. in 1991. Thus, there is the potential for selection on resistance to these two herbivores. Despite the many significant phenotypic correlations between herbivore abundances within a year, most genetic correlations between herbivore abundances within a year were not significant. The genetic and phenotypic correlation structure varied from year to year in this three-year study. Thus, it appears that there is the potential for evolution of resistance to the two herbivores for which we found significant heritabilities, but multispecies coevolution seems unlikely.     

Tolerance to deer herbivory and resistance to insect herbivores in the common evening primrose (Oenothera biennis)

The evolution of plant defence in response to herbivory will depend on the fitness effects of damage, availability of genetic variation and potential ecological and genetic constraints on defence. Here, we examine the potential for evolution of tolerance to deer herbivory in Oenothera biennis while simultaneously considering resistance to natural insect herbivores. We examined (i) the effects of deer damage on fitness, (ii) the presence of genetic variation in tolerance and resistance, (iii) selection on tolerance, (iv) genetic correlations with resistance that could constrain evolution of tolerance and (v) plant traits that might predict defence. In a field experiment, we simulated deer damage occurring early and late in the season, recorded arthropod abundances, flowering phenology and measured growth rate and lifetime reproduction. Our study showed that deer herbivory has a negative effect on fitness, with effects being more pronounced for late‐season damage. Selection acted to increase tolerance to deer damage, yet there was low and nonsignificant genetic variation in this trait. In contrast, there was substantial genetic variation in resistance to insect herbivores. Resistance was genetically uncorrelated with tolerance, whereas positive genetic correlations in resistance to insect herbivores suggest there exists diffuse selection on resistance traits. In addition, growth rate and flowering time did not predict variation in tolerance, but flowering phenology was genetically correlated with resistance. Our results suggest that deer damage has the potential to exert selection because browsing reduces plant fitness, but limited standing genetic variation in tolerance is expected to constrain adaptive evolution in O. biennis.     

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.     

Association mapping of plant resistance to insects

Association mapping is rapidly becoming an important method to explore the genetic architecture of complex traits in plants and offers unique opportunities for studying resistance to insect herbivores. Recent studies indicate that there is a trade-off between resistance against generalist and specialist insects. Most studies, however, use a targeted approach that will easily miss important components of insect resistance. Genome-wide association mapping provides a comprehensive approach to explore the whole array of plant defense mechanisms in the context of the generalist-specialist paradigm. As association mapping involves the screening of large numbers of plant lines, specific and accurate high-throughput phenotyping (HTP) methods are needed. Here, we discuss the prospects of association mapping for insect resistance and HTP requirements.     

Testing genotypic variation of an invasive plant species in response to soil disturbance and herbivory

Herbivores, competitors, and predators can inhibit biological invasions (“biotic resistance” sensu Elton 1959), while disturbance typically promotes biological invasions. Although biotic resistance and disturbance are often considered separately in the invasion literature, these two forces may be linked. One mechanism by which disturbance may facilitate biological invasions is by decreasing the effectiveness of biotic resistance. The effects of both disturbance and biotic resistance may vary across invading genotypes, and genetic variation in the invasive propagule pool may increase the likelihood that some genotypes can overcome biotic resistance or take greater advantage of disturbance. We conducted an experimental field trial in which we manipulated soil disturbance (thatch removal and loosening soil) and the presence of insect herbivores and examined their effects on the invasion success of 44 Medicago polymorpha genotypes. As expected, insecticide reduced leaf damage and increased Medicago fecundity, suggesting that insect herbivores in this system provide some biotic resistance. Soil disturbance increased Medicago fecundity, but did not alter the effectiveness of biotic resistance by insect herbivores. We found significant genetic variation in Medicago in response to disturbance, but not in response to insect herbivores. These results suggest that the ability of Medicago to invade particular habitats depends on the amount of insect herbivory, the history of disturbance in the habitat, and how the specific genotypes in the invader pool respond to these factors.     

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.     

Heritable variation in the foliar secondary metabolite sideroxylonal in <Emphasis Type="Italic">Eucalyptus</Emphasis> confers cross-resistance to herbivores

Plants encounter a broad range of natural enemies and defend themselves in diverse ways. The cost of defense can be reduced if a plant secondary metabolite confers resistance to multiple herbivores. However, there are few examples of positively correlated defenses in plants against herbivores of different types. We present evidence that a genetically variable chemical trait that acts as a strong antifeedant to mammalian herbivores of Eucalyptus also deters insect herbivores, suggesting a possible mechanism for cross-resistance. We provide field confirmation that sideroxylonal, an important antifeedant for mammalian herbivores, also determines patterns of damage by Christmas beetles, a specialist insect herbivore of Eucalyptus. In a genetic progeny trial of Eucalyptus tricarpa, we found significant heritabilities of sideroxylonal concentration (0.60), overall insect damage (0.34), and growth traits (0.30–0.53). Population of origin also had a strong effect on each trait. Negative phenotypic correlations were observed between sideroxylonal and damage, and between damage and growth. No relationship was observed between sideroxylonal concentration and any growth trait. Our results suggest that potential for evolution by natural selection of sideroxylonal concentrations is not strongly constrained by growth costs and that both growth and defense traits can be successfully incorporated into breeding programs for plantation trees.     

The constraints of selecting for insect resistance in plantation trees

• 1 High productivity in plantations of exotic tree species is achieved by management for fast growth in the absence of the full complex of co‐evolved insect herbivores. In the case of Eucalyptus, silvicultural selection for desirable wood traits is concomitant with a trade‐off against defence and a reduction of chemical and genetic diversity. These factors, combined with accidental introductions, rapid insect evolution and the emergence of new pests, increase the likelihood that future plantations will need insect pest management to maintain productivity.
• 2 Forestry researchers have suggested that selecting for resistant genotypes may be beneficial in insect control. There are, however, significant differences between long‐lived trees and annual crops that make this approach unlikely to be successful. This is illustrated using several examples of research into resistance to insect herbivores in trees.
• 3 Selection for resistance to insects in trees requires an assessment of trial plantations for heritable variation in insect damage and then a determination of the effect of variation in resistance on insect population parameters. Identifying rare resistant genotypes using markers is difficult because many factors interact to produce a resistant phenotype, and phytophagous insects have less intimate relationships with their host than pathogens, resulting in weak associations with genetic loci.
• 4 If resistant genotypes are identified, their widespread deployment in plantations might not provide satisfactory management of insect pests when the use of extensive monocultures is continued. In this paper, experiments are suggested that would explore the effectiveness of polycultures or chemotype mixtures with respect to ameliorating the damage of insects on plantation productivity. In addition, mitigating the effects of some insects on plantation productivity by maintaining vigour of fast‐growing eucalypts should be considered.

     

Effects of drought, temperature, herbivory, and genotype on plant–insect interactions in soybean (Glycine max)

Climate change is predicted to cause continued increases in global temperatures, greater variability in precipitation and in some cases, more frequent insect pest outbreaks. Here we seek to understand how abiotic and biotic stresses associated with climate change can affect plant-herbivore interactions in a model crop species (soybean, Glycine max (L.) Merr.) by answering three questions: (1) Do the combined effects of abiotic and biotic stresses associated with climate change cause synergistic negative effects on plant biomass? (2) Can abiotic stress affect resistance of plants to insect herbivores? (3) Does genetic variation in plant traits modify a plant’s response to stress? We performed three experiments in controlled growth environments using up to 51 soybean genotypes selected to vary in numerous traits associated with drought and resistance against pests (e.g., insect herbivores, nematodes, and pathogenic fungi), and up to 3 generalist-feeding herbivorous noctuid moth species (Helicoverpa zea, Heliothis virescens, and Spodoptera exigua) that commonly feed on soybean in North America. Drought and herbivory had the largest and the most consistent negative effects on plant performance, reducing the above- and below-ground biomass by 10-45 %, whereas increased temperature had little to no effect on plants. Drought also increased susceptibility to generalist noctuid herbivores, but these results varied dramatically in magnitude and direction among plant genotypes. Our experiments show that the effects of abiotic and biotic stress on soybean biomass were largely due to the additive effects of these stresses, and there exists substantial genetic variation in the soybean germplasm pool we studied that could be used as a source of parental stock in breeding new crops that can more effectively tolerate and resist the combined negative effects of insect herbivory and drought.     

Specialist Insect Herbivore and Light Availability Do Not Interact in the Evolution of an Invasive Plant

Release from specialist insect herbivores may allow invasive plants to evolve traits associated with decreased resistance and increased competitive ability. Given that there may be genetic trade-off between resistance and tolerance, invasive plants could also become more tolerant to herbivores. Although it is widely acknowledged that light availability affects tolerance to herbivores, little information is available for whether the effect of light availability on tolerance differ between the introduced and native populations. We conducted a common garden experiment in the introduced range of Alternanthera philoxeroides using ten invasive US and ten native Argentinean populations at two levels of light availability and in the presence or absence of a specialist stem-boring insect Agasicles hygrophila. Plant biomass (total and storage root biomass), two allocation traits (root/shoot ratio and branch intensity, branches biomass/main stem biomass) and two functional traits (specific stem length and specific leaf area), which are potentially associated with herbivore resistance and light capture, were measured. Overall, we found that A. philoxeroides from introduced ranges had comparable biomass and tolerance to specialist herbivores, lower branch intensity, lower specific stem length and specific leaf area. Moreover, introduced populations displayed higher shade tolerance of storage root biomass and lower plastic response to shading in specific stem length. Finally, light availability had no significant effect on evolution of tolerance to specialist herbivores of A. philoxeroides. Our results suggest that post-introduction evolution might have occurred in A. philoxeroides. While light availability did not influence the evolution of tolerance to specialist herbivores, increased shade tolerance and release from specialist insects might have contributed to the successful invasion of A. philoxeroides.     

Gene flow and geographically structured coevolution

Many of the dynamic properties of coevolution may occur at the level of interacting populations, with local adaptation acting as a force of diversification, as migration between populations homogenizes these isolated interactions. This interplay between local adaptation and migration may be particularly important in structuring interactions that vary from mutualism to antagonism across the range of an interacting set of species, such as those between some plants and their insect herbivores, mammals and trypanosome parasites, and bacteria and plasmids that confer antibiotic resistance. Here we present a simple geographically structured genetic model of a coevolutionary interaction that varies between mutualism and antagonism among communities linked by migration. Inclusion of geographic structure with gene flow alters the outcomes of local interactions and allows the maintenance of allelic polymorphism across all communities under a range of selection intensities and rates of migration. Furthermore, inclusion of geographic structure with gene flow allows fixed mutualisms to be evolutionarily stable within both communities, even when selection on the interaction is antagonistic within one community. Moreover, the model demonstrates that the inclusion of geographic structure with gene flow may lead to considerable local maladaptation and trait mismatching as predicted by the geographic mosaic theory of coevolution.     

Natural selection and the joint evolution of toleranceand resistance as plant defenses

Plants can defend themselves against the damaging effects of herbivory in at least two ways. Resistant plants avoid or deter herbivores and are therefore fed upon less than susceptible plants. Tolerant plants are not eaten less than plants with little tolerance, but the effects of herbivore damage are not so detrimental to a tolerant plant as they are to a less tolerant plant. Biologists have suggested that these two strategies might represent two alternative and redundant defenses against herbivory since they appear to serve the same function for plants. I explore the relationship between resistance and tolerance, particularly with regards to how the joint evolution of these two traits will influence the evolution of plant defense. Although I briefly review some of the contributions of theory to the study of tolerance, I concentrate on an empirical, ecological genetic approach to the study of the evolution of these characters and the coevolution of tolerance and herbivores. In order to understand the evolution of any trait, we must understand the evolutionary forces acting on the trait. Specifically, we must understand how natural selection acts on tolerance. I review several studies that have specifically measured the form of selection acting on tolerance and tested the hypothesis that resistance and tolerance are alternative strategies. I also present a statistical analysis that does not support the hypothesis that herbivores are selective agents on tolerance. Finally, I consider a variety of constraints that possibly restrict the evolution of tolerance. This revised version was published online in July 2006 with corrections to the Cover Date.     

Genetic architecture of resistance to aphids and mites in a willow hybrid system

Hybrid plants often differ in resistance to arthropods compared to the parental species from which they are derived. To better understand the relative contribution of genetic effects in influencing plant resistance to arthropods, we examined the genetic architecture of resistance in a willow hybrid system, Salix eriocephala, S. sericea, and their interspecific hybrids. Resistance to two arthropods, a willow leaf aphid (Chaitophorus sp.: Aphididae) and an eriophyoid mite (Aculops tetanothrix: Eriophyidae), were compared because resistance to different herbivores may be controlled by different traits and influenced by different genetic effects. We found additive and nonadditive genetic effects to be important in explaining the difference between willow species in resistance to aphids and mites. F2 hybrids exhibited low resistance to aphids, suggesting breakdown of favourable epistatic interactions that confer resistance. F2 hybrids, however, exhibited high resistance to mites, suggesting either the breakdown of interactions that affect traits used by mites in host location or the creation of favourable epistatic interactions. This study demonstrates the potential role of herbivores in affecting plant genetic structure, such that selection by herbivores can potentially lead to the creation of gene interactions that influence host resistance traits or host recognition traits used by the herbivore.     

Phytochemical defences and performance of specialist and generalist herbivores: a meta-analysis

1. Phytochemical coevolution theory, a long-standing paradigm in plant–insect interactions, predicts that specialist herbivores are less negatively affected by the allelochemicals of their host plants than are generalist herbivores. Although this theory is prevalent in plant–insect science, it is not always supported by empirical studies measuring the performance of specialist and generalist insects in response to allelochemicals. 2. The present study aimed to investigate: (i) whether there a difference between specialist and generalist performance in response to allelochemicals and (ii) whether the effect of allelochemicals on specialists and generalists depend upon allelochemical class or insect order. 3. A meta-analysis was conducted incorporating 76 effect sizes drawn from studies that directly compared the performance of specialist and generalist insects in response to treatment and control diets. Most of the effect sizes were related to the performance metric growth, the insect order Lepidoptera, and the allelochemical class nitrogen-containing compounds. 4. As predicted by phytochemical coevolution theory, specialist insects responded less negatively to allelochemicals of their hosts than generalist insects in terms of growth. There were no significant differences in terms of fecundity or survival, or among allelochemical classes or insect orders. 5. These results support the prediction of phytochemical coevolution theory that specialist insects respond less negatively to allelochemicals of their hosts than generalists, although only in terms of growth.     

欧洲山芥种质资源的表型遗传多样性分析

欧洲山芥是研究抗虫皂苷生物合成及植食昆虫与植物协同进化机制的理想模式植物,是极具开发潜力的十字花科植物。为加强欧洲山芥种质资源的研究利用,对保存于国家蔬菜种质资源中期库中引自欧洲的33份欧洲山芥种质和1份杂交材料进行了物候期观测、主要形态学性状测定和小菜蛾抗性鉴定。结果表明:多数材料的表型性状的差异较大,材料间现蕾期最早(BV13-BAR13)和最晚(BV13-BAR9)的2份材料相差达47 d;从始花期到末花期的天数分布在17~46 d之间,总体上花期较短;叶表光滑的种质居多;还发现4份角果抱茎的种质;极差和变异系数最大的性状是株高、角果长和花序长。34份材料中仅9份感小菜蛾,其他均具有很高的田间抗性。基于所有表型性状的聚类分析将34份欧洲山芥分为3个类群,抗虫和感虫材料分属不同的类群(F1、BV13-BAR8和BV13-BAR22除外),基本反映出各类群的亲缘关系。本研究结果将为欧洲山芥的遗传改良以及欧洲山芥种质资源在十字花科作物生产和育种上的应用提供基本依据。