GENETICS OF BRASSICA RAPA. 3. COSTS OF DISEASE RESISTANCE TO THREE FUNGAL PATHOGENS

Genetic costs of resistance to pathogens may be an important factor maintaining heritable variation for resistance in natural populations. Pleiotropic fitness trade-offs occur when genetic resistance causes reduction in other components of fitness. Although costs of resistance have an important influence on plant-pathogen interactions, few previous studies have detected pleiotropic costs of resistance in the absence of confounding effects of linkage disequilibrium. To avoid this potential problem, we performed artificial selection experiments on resistance to two fungal pathogens, Leptosphaeria maculans, and Peronospora parasitica, and compared growth rates of resistant and susceptible genotypes of Brassica rapa in the absence of pathogens. Leptosphaeria resistance had no effect on growth rate, indicating cost-free defense. In contrast, Peronospora-resistant genotypes grow 6% slower than Peronospora-susceptible genotypes in pathogen-free environments, indicating a significant genetic fitness cost to Peronospora resistance. Such genetic trade-offs could maintain genetic variation in the wild. Another factor that might explain heritable variation for resistance is ecological trade-offs, in which genetic resistance to one species causes susceptibility to another. Such ecological trade-offs do not exist for the pathogens studied in this system.     

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.     

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.     

EXPERIMENTAL EVOLUTION OF RESISTANCE IN BRASSICA RAPA: CORRELATED RESPONSE OF TOLERANCE IN LINES SELECTED FOR GLUCOSINOLATE CONTENT

The evolutionary response of plant populations to selection for increased defense may be constrained by costs of defense. The purpose of this study was to investigate such constraints on the evolution of defense due to a cost of defense manifested as a trade-off between defense and tolerance. Variation in the response to artificial damage (tolerance) among lines of Brassica rapa that had been artificially selected for foliar glucosinolate content (defense) was examined. Leaf area was removed from replicates of three selection lines (high glucosinolates, control, and low glucosinolates) at three damage levels (0%, 20%, and 60% damage). An external cost of defense would result in a statistically significant selection line by damage treatment interaction, with those selected for high defense expressing less tolerance than those selected for low defense. Damage treatment had a significant overall effect on estimated total fitness, with fitness declining with increasing damage level. Further, selection line also had a significant overall effect on estimated total fitness, with low-defense selection lines having higher fitness compared to both control and high-defense selection lines. More importantly, a cost of defense in terms of tolerance was demonstrated by a significant selection line-by-damage treatment interaction. This interaction was in the direction to demonstrate a genetic trade-off between defense and tolerance, with low-defense selection lines decreasing estimated total fitness in response to damage less than both control and high-defense selection lines. Variation in tolerance among selection lines was due to the greater ability of low-defense lines to maintain fruit and seed production despite the presence of damage. In terms of tolerance, this cost of glucosinolate production in B. rapa could constrain the evolution of increased defense and, in so doing, maintain individuals within the population that are poorly defended yet tolerant.     

COSTS OF COUNTERDEFENSES TO HOST RESISTANCE IN A PARASITOID OF DROSOPHILA

Abstract The ability of a parasitoid to evolve enhanced counterdefenses against host resistance and its possible costs were studied in a Drosophila -parasitoid system. We reared Asobara tabida (Braconidae, Hymenoptera) exclusively on D. melanogaster to impose artificial selection for improved counterdefenses against cellular encapsulation, the main host defense against parasitism. Controls were reared on D. subobscura , the main host of the population of wasps from which the laboratory culture was derived and a species that never encapsulates parasitoids. We observed improved survival and avoidance of encapsulation in all five selection lines compared to their paired control lines, although there was unexpected variation among pairs. Improved survival was associated with parasitoid eggs becoming embedded in host tissue, where they were protected from circulating haemocytes. There were no differences among lines in average adult size, fat content, egg load, or performance on D. subobscura . However, the duration of the egg stage in selection lines was longer than that of control lines, probably because of reduced nutrient and/or oxygen supply when eggs are embedded in host tissue. We suggest that this delay in hatching reduces the probability of parasitoid survival if another parasitoid egg is laid in the same host (superparasitism or multiparasitism) and hence is a cost of enhanced counterdefenses against host resistance.     

线性规划法在植物生态学数学模型参数估计中的应用初探

本文提出了一种植物生态学数学模型参数估计的新方法——线性规划法,并结合最小二乘法对模型参数的估计进行了实例分析。认为某种程度上前者更优于后者。是一种值得进一步研究的方法。     

SEX-SPECIFIC COSTS OF RESISTANCE TO THE FUNGAL PATHOGEN USTILAGO VIOLACEA (MICROBOTRYUM VIOLACEUM) IN SILENE ALBA

Costs of resistance are often invoked to explain the maintenance of polymorphisms for resistance to fungal pathogens in natural plant populations. To investigate such costs, 27 half-sib families of Silene alba, collected from a single host population, were grown in experimental populations in the presence and absence of the anther-smut fungus Ustilago violacea, a host-sterilizing pathogen transmitted by insects that are both pollinators and vectors of the disease. Host families differed significantly in resistance to inoculation, indicating the presence of genetic variation for mechanisms that impede fungal growth once the disease is encountered (“biochemical” resistance) within the host population. In addition, host families differed significantly in onset of flowering and in flower production in the absence of the disease. Path analysis revealed that late onset of flowering in male host families made a direct contribution to high field resistance (P < 0.01), probably due to a reduced rate of contact between hosts and vectors carrying high spore loads (avoidance, or “phenological” resistance). The contribution of low flower production to field resistance only approached significance (P < 0.10). There was a significantly positive genetic association between biochemical and phenological resistance, suggesting that delayed flowering is either a pleiotropic effect of biochemical resistance, or that genes governing these traits are in linkage disequilibrium. Path analysis revealed that biochemical resistance made both a direct contribution to field resistance (P < 0.01) and a positive indirect contribution via its association with phenology and flower production (P < 0.05) in male hosts. Costs of resistance were sex specific. Male host families with high field resistance had significantly lower reproductive success in healthy populations, indicating a fitness cost of field resistance (P < 0.01), whereas no costs were detected for female hosts. Path analysis revealed that the biochemical component of field resistance made no direct contribution to the observed fitness cost in male hosts, whereas its indirect effect through phenology was only marginally significant (P < 0.10). This finding indicates that fitness costs were mainly due to the phenological component of field resistance. Because the host population had no known history of disease, it is not clear whether the fitness costs are responsible for maintenance of the resistance polymorphism or whether the polymorphism is present for reasons unrelated to pathogen infection. Interactions between host families and pathogen strains with respect to inoculation success were not significant. Hence, there was no evidence for indirect costs of biochemical resistance, that is, reduced resistance to alternative strains. Infection rates in experimental populations with an initially patchy distribution of the pathogen were lower than in populations with a uniform pathogen distribution, suggesting that the effective pathogen pressure and hence the relative success of susceptible and resistant individuals may, in addition to fitness costs of resistance, depend on the spatial population structure of the pathogen.     

半干旱生态条件下植物个体的综合生态效应的空间距离分布规律

通过对有关生态因子的空间距离变化规律的测定和引入相应的生态效应系数综合分析,得到在半干旱生态条件下植物个体的综合生态效应的空间距离分布规律为: E=α_1E_(1o)[1 (x/hc)~2]-3/2 (α_NE_(NO) α_WE_(WO))[1 (x/hr)~2]-3/2 (α_cE_(co) α_hE_(ho))[1 (x/hc)~2]-1/2由于具抑制效应的因子的生态效应随相对空间距离增加而减弱的速度大于其互惠效应的因子,在一定相对空间距离处(w/h=6.8)综合生态效应由干扰转变为互惠。定义了生态场的概念。     

COSTS AND BENEFITS OF PLANT RESPONSES TO DISEASE: RESISTANCE AND TOLERANCE

A major assumption of models of the evolution of plant resistance to disease is that plant resistance involves fitness costs. To test this assumption, a field experiment was performed so that a quantitative-genetic analysis could be used to detect fitness costs to Ipomoea purpurea of resistance to different fungal isolates of Colletotrichum dematium, a pathogenic fungus causing the disease anthracnose. This experiment yielded no evidence that resistance to anthracnose involves direct fitness costs. Nevertheless, trade-offs in plant fitness that were unrelated to resistance were detected between different disease environments. Tolerance, defined as the ability to compensate in part for fitness decrements caused by disease, was found to involve fitness costs. Halfsib families that were more tolerant of disease had lower fitness in the absence of disease. The possibility that the cost of tolerance could obscure fitness costs of resistance is explored.     

潮滩红树植物抗低温适应的生态学研究

采用电导法定量测定了中国东南沿海红树植物叶片的抗寒力变化。结果表明;(1)海南琼山东寨港主要红树种类冬季抗寒力(半致死温度)介于-2.3～-6.8℃之间,其中红树科植物抗寒力较强,多数种类属于低温敏感性相对较弱的L类型;海桑科和楝科红树种类抗寒力较弱,属于低温敏感性较强的H类型。(2)随纬度升高(个别地点除外),秋茄(Kandelia candel)、桐花树(Aegiceras corniculatum)和红海榄(Rhizophors,stylosa)抗寒力增强,呈现种群分化趋势。(3)同一地点,秋茄和桐花树抗寒力有明显季节变化：夏季最低,秋冬两季高于春季。(4)同一群落内,生长于中潮区滩面的秋茄抗寒力强于高潮区,但桐花树抗寒力在各个滩面相差木大。(5)秋茄和桐花树树冠上不同部位叶片的抗寒力不同,由表及里,由上至下,呈降低趋势。红树叶片抗寒力的种群分化、季节变化和部位差异是其趋异适应的结果,有益于红树植物的生存和繁育。     

梯度分析技术用于植物抗旱性的研究

将梯度分析技术引入植物抗旱性的研究,对植物的解剖结构指标进行分析。在ＰＣＡ分析结果的基础,通过引进ＰＣ间的因子ａｊ,梯度分析技术用于植物抗旱性的研究比用于植物群落的研究,其结果更具有观性,明确性,是研究植物抗旱性的一个可行、有效的技术手段。     

四种植物种子萌发及苗期抗旱性差异的研究

对四种豆科植物种子萌发及苗期抗旱性作了比较研究,结果表示：种子萌发期抗旱笥强的植物其苗期抗旱性亦强。干旱引起组织脱水,植株生长受到抑制,幼苗出现衩始萎蔫时间与细胞膜受害时间一致。四种植物显示萎蔫时土壤含水量明显不同,红豆草为４．８０％、羊柴２．２％、花棒１．７４％、柠条１．５１％．据种子相对发芽率、膜透性变化等生理指标综合评定四种植物种子萌发及苗期抗旱能力次序为：柠条＞花棒＞羊柴＞红豆草。     

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.     

中国珍贵植物药生态分布研究

本文概述珍贵植物药在中国三大自然区域分布的一般生态学特性,探讨了地域性分布规律及生长地对药材质量,疗效的影响,研究表明：南方山势峻峭,气候温暖多变,植物药资源种类多于北方;北方山体高大,山势平缓,气候寒冷,多干燥,植物群落单一,蕴藏量多于南方。     

EPISTATIC INTERACTIONS CAN LOWER THE COST OF RESISTANCE TO MULTIPLE CONSUMERS

It is widely assumed that resistance to consumers (e.g., predators or pathogens) comes at a “cost,” that is, when the consumer is absent the resistant organisms are less fit than their susceptible counterparts. It is unclear what factors determine this cost. We demonstrate that epistasis between genes that confer resistance to two different consumers can alter the cost of resistance. We used as a model system the bacterium Escherichia coli and two different viruses (bacteriophages), T4 and Λ, that prey upon E. coli. Epistasis tended to reduce the costs of multiple resistance in this system. However, the extent of cost savings and its statistical significance depended on the environment in which fitness was measured, whether the null hypothesis for gene interaction was additive or multiplicative, and subtle differences among mutations that conferred the same resistance phenotype.     

THE ROLE OF PHOSPHOTASE AND ACETYLCHOLINESTERASE IN RESISTANCE TO PHOXIM IN HELICOVERPA ARMIGERA

Abstract The resistance mechanism underlying phoxim resistance was investigated through biochemical assays. The results show that there are marked relations between the phosphatase activity and resistance to phoxim. Furthermore, there is a significant difference in the K _i values of AChE between the two strains and it is inferred that the change in ability of binding to phoxim is a mechanism responsible for resistance to phoxim. The metabolic inhibitors Triphenyl Phosphate (TPP) and O,O-diethyl-O-phenyl-thiophosphate (SV₁) displayed significant synergism of phoxim toxicity to the two strains. It implied that esterase might be involved in resistance. However, there is no marked correlation between resistance and CarE.     

GENETIC BASIS OF A BETWEEN-ENVIRONMENT TRADE-OFF INVOLVING RESISTANCE TO CADMIUM IN DROSOPHILA MELANOGASTER

In a replicated, laboratory, natural selection experiment Drosophila melanogaster populations were maintained for 20 generations either on unpolluted medium or on polluted medium containing cadmium chloride at a concentration of 80 μg/ml. Lines maintained on polluted medium evolved resistance. In comparison with unpolluted lines, their juvenile survivorship increased from 35% to 46%, developmental period decreased from 13.7 days to 13.0 days, and fecundity increased from 3 to 29 eggs per two-day period. Emergence weights, however, did not change. By contrast the “environmental” effect of moving susceptible flies onto polluted medium was that after two generations survivorship fell 62%, developmental period increased 40%, and fecundity fell 97%. Emergence weights fell 31% in females and 28% in males. Resistant lines paid a fitness cost in unpolluted environments, with fecundity being reduced by 44% and emergence weights being reduced by 4% in females and 6% in males. Developmental period, however, was unaffected. Analyses of crosses and backcrosses between the lines suggested that the evolved cadmium resistance was due to a single sex-linked gene. Levels of dominance were calculated, and in each life-history character the resistant allele was found to be completely dominant. Because the life-history effects appear to be produced by a single gene, it is probable that they all depend on the same metabolic pathway. Metallothionein production is a likely candidate because this is known to be controlled by genes on the X-chromosome. The study adds to a small number of examples of single or closely linked genes with large antagonistic pleiotropic effects on life histories. The result here is a between-environment trade-off, allowing animals increased fitness in polluted environments, but only at the cost of reduced growth and reproduction in unpolluted environments.     

THE DISTRIBUTION AND ORIGIN OF GENES FOR RACE-SPECIFIC RESISTANCE TO MELAMPSORA LINI IN LINUM MARGINALE

The genetic basis of resistance in wild flax (Linum marginale) to its host-specific pathogen Melampsora lini was investigated in seven lines collected from a single population growing at Kiandra, New South Wales and in an additional ten lines collected more widely across southeastern Australia. All lines showed different phenotypic patterns of resistance and susceptibility. Genetic analyses indicated the presence of single dominant genes for race-specific resistance in all but one of these lines. That particular line appeared to carry two linked dominant genes for resistance. Intercrosses between lines in each of these groups of L. marginale detected substantially more linkage between the resistance genes in the Kiandra population sample than between those in the broader geographic collection. This result is interpreted to indicate a possible mechanism whereby resistance genes are generated in natural populations.