Aggressiveness is associated with genetic diversity in landlocked salmon (Salmo salar)

The amount of intraindividual genetic variation has often been found to have profound effects on life history traits. However, studies concerning the relationship between behaviour and genetic diversity are scarce. Aggressiveness is an important component of competitive ability in juvenile salmonids affecting their later performance and survival. In this study, we used an experimental approach to test the prediction that juveniles with low estimated genetic diversity should be less aggressive than juveniles with high estimated genetic diversity in fry from a highly endangered population of land-locked salmon (Salmo salar). This was achieved by using a method enabling the accurate estimation of offspring genetic diversity based on parental microsatellite genotype data. This allowed us to create two groups of offspring expected to have high or low genetic diversity in which aggressive behaviour could be compared. Salmon fry with low estimated genetic diversity were significantly less aggressive than fry with high estimated genetic diversity. Closer analysis of the data suggested that this difference was due to differences in more costly acts of aggression. Our result may reflect a direct effect of genetic variation on a fitness-related trait; however, we cannot rule out an alternative explanation of allele-specific phenotype matching, where lowered aggression is expressed towards genetically more similar individuals.     

Delayed Phenotypic Expression of Growth Hormone Transgenesis during Early Ontogeny in Atlantic Salmon (Salmo salar)?

Should growth hormone (GH) transgenic Atlantic salmon escape, there may be the potential for ecological and genetic impacts on wild populations. This study compared the developmental rate and respiratory metabolism of GH transgenic and non-transgenic full sibling Atlantic salmon during early ontogeny; a life history period of intense selection that may provide critical insight into the fitness consequences of escaped transgenics. Transgenesis did not affect the routine oxygen consumption of eyed embryos, newly hatched larvae or first-feeding juveniles. Moreover, the timing of early life history events was similar, with transgenic fish hatching less than one day earlier, on average, than their non-transgenic siblings. As the start of exogenous feeding neared, however, transgenic fish were somewhat developmentally behind, having more unused yolk and being slightly smaller than their non-transgenic siblings. Although such differences were found between transgenic and non-transgenic siblings, family differences were more important in explaining phenotypic variation. These findings suggest that biologically significant differences in fitness-related traits between GH transgenic and non-transgenic Atlantic salmon were less than family differences during the earliest life stages. The implications of these results are discussed in light of the ecological risk assessment of genetically modified animals.     

Elevated ability to compete for limited food resources by 'all-fish' growth hormone transgenic common carp Cyprinus carpio

Food consumption, number of movements and feeding hierarchy of juvenile transgenic common carp Cyprinus carpio and their size-matched non-transgenic conspecifics were measured under conditions of limited food supply. Transgenic fish exhibited 73·3% more movements as well as a higher feeding order, and consumed 1·86 times as many food pellets as their non-transgenic counterparts. After the 10 day experiment, transgenic C. carpio had still not realized their higher growth potential, which may be partly explained by the higher frequency of movements of transgenics and the 'sneaky' feeding strategy used by the non-transgenics. The results indicate that these transgenic fish possess an elevated ability to compete for limited food resources, which could be advantageous after an escape into the wild. It may be that other factors in the natural environment ( i.e. predation risk and food distribution), however, would offset this advantage. Thus, these results need to be assessed with caution.     

Food consumption and growth of Atlantic salmon Salmo salar parr in sub-Arctic rivers: empirical support for food limitation and competition

The quantitative food consumption and somatic growth of Atlantic salmon Salmo salar parr were compared between three sub-Arctic rivers in northern Norway and Finland, addressing the potential occurrence of resource limitation and interspecific competition. In one of the rivers, previous resource partitioning studies have suggested severe dietary competition between juvenile S. salar and a dense population of alpine bullheads Cottus poecilopus . It was hypothesized that S. salar parr in this river would have restricted food consumption and growth rates compared to the S. salar populations in the other two rivers where interspecific competition was less likely to occur. The feeding and growth performance differed significantly between the S. salar populations. The lowest food acquisition and growth rates were in the S. salar parr population living in sympatry with C. poecilopus , confirming a restricted food supply for the S. salar parr and providing empirical support for the presence of resource limitation and interspecific food competition in this river system. The study reveals that S. salar parr in sub-Arctic rivers may experience food limitations resulting in diminished growth rates.     

食物供应量对转红色荧光蛋白基因唐鱼 生存和生长的影响

选取雄性杂合子转红色荧光蛋白基因唐鱼(Tanichthys albonubes)(简称:转基因唐鱼)与雌性非转基因唐鱼交配产卵,出膜7d后,在水温(25.0 ±2.0)℃条件下,选择健康仔鱼进行饱食(食物供应量高)、半饥饿(食物供应量中)和饥俄(食物供应量低)3种处理,结果显示:(1)出膜后7～72 d的唐鱼饱食组与半...     

Early Life-History Consequences of Growth-Hormone Transgenesis in Rainbow Trout Reared in Stream Ecosystem Mesocosms

There is persistent commercial interest in the use of growth modified fishes for shortening production cycles and increasing overall food production, but there is concern over the potential impact that transgenic fishes might have if ever released into nature. To explore the ecological consequences of transgenic fish, we performed two experiments in which the early growth and survival of growth-hormone transgenic rainbow trout (Oncorhynchus mykiss) were assessed in naturalized stream mesocosms that either contained predators or were predator-free. We paid special attention to the survival bottleneck that occurs during the early life-history of salmonids, and conducted experiments at two age classes (first-feeding fry and 60 days post-first-feeding) that lie on either side of the bottleneck. In the late summer, the first-feeding transgenic trout could not match the growth potential of their wild-type siblings when reared in a hydrodynamically complex and oligotrophic environment, irrespective of predation pressure. Furthermore, overall survival of transgenic fry was lower than in wild-type (transgenic = 30% without predators, 8% with predators; wild-type = 81% without predators, 31% with predators). In the experiment with 60-day old fry, we explored the effects of the transgene in different genetic backgrounds (wild versus domesticated). We found no difference in overwinter survival but significantly higher growth by transgenic trout, irrespective of genetic background. We conclude that the high mortality of GH-transgenic trout during first-feeding reflects an inability to sustain the basic metabolic requirements necessary for life in complex, stream environments. However, when older, GH-transgenic fish display a competitive advantage over wild-type fry, and show greater growth and equal survival as wild-type. These results demonstrate how developmental age and time of year can influence the response of genotypes to environmental conditions. We therefore urge caution when extrapolating the results of GH-transgenesis risk assessment studies across multiple life-history or developmental stages.     

Increased intrinsic growth rate is advantageous even under ecologically stressful conditions in coho salmon (<Emphasis Type="Italic">Oncorhynchus kisutch</Emphasis>)

Growth rate is an ecologically important trait, affecting the energy acquisition from, and provisioning to, the surrounding community. One of many costs suggested to counteract the evolution of increased intrinsic growth rate is an associated reduction in tolerance to conditions of nutrient stress. Here we test this concept with individuals possessing experimentally increased intrinsic growth rates (growth hormone transgenic coho salmon, Oncorhynchus kisutch) relative to wild genotypes. Using a series of three experiments, survival and growth of both genotypes were assessed on a physiological and behavioral level while varying food abundance, social interactions, and predation risk. Only in complete absence of exogenous food in newly emerged fry did the high intrinsic growth rate appear costly with a shorter average survival time compared to wild-type (Exp. 1). In experiment 2, genotypes with elevated intrinsic growth showed equal or higher survival and growth than wild-type genotypes In a third experiment, adding very limited amounts of food and allowing for social interactions in a simulated natural environment benefited transgenic individuals relative to wild-types, but at similar magnitudes in both the absence and presence of predators. Populations with transgenic individuals present did not crash under these competitive conditions as previously reported when studied in simple environments where hiding and attack escape were not possible. Our data suggest that transgenic fish have a greater scope for growth under most conditions, but are not obligated to use this capability. Physiological (e.g. appetite and conversion efficiency) and behavioral traits (e.g. competitive ability and risk-taking) found previously to correlate positively with intrinsic growth rate in the transgenic strain likely aided in their survival and growth, even under food limited conditions. Hence, at least in coho salmon, intrinsic growth rate does not appear to strongly affect survival under nutrient stress.     

Selection on increased intrinsic growth rates in coho salmon, Oncorhynchus kisutch

Substantial evidence from the animal kingdom shows that there is a trade-off between benefits and costs associated with rapid somatic growth. One would therefore expect growth rates under natural conditions to be close to an evolutionary optimum. Nevertheless, natural selection in many salmonid species appears to be toward larger size and earlier emergence from spawning redds, indicating a potential for increased growth rate to evolve. We tested how selection for genetic variants (growth hormone transgenic coho salmon, Oncorhynchus kisutch, with more than doubled daily growth rate potential relative to wild genotypes) depended on predator timing and food abundance during the early period of life (fry stage). In artificial redds, fry of the fast-growing genotypes showed a highly significant developmental shift, emerging from gravel nests approximately two weeks sooner, but with an 18.6% reduced survival, relative to wild-genotype fry. In seminatural streams, fry of the fast-growing genotypes suffered higher predation than those of wild genotypes when predators were present at the time of fry emergence, but this difference was less pronounced when food was scarce. In streams where predators were introduced after emergence, fry survived equally well regardless of food availability. Surviving fry grew faster in habitats provided with more food, and fast-growing genotypes also grew faster than wild genotypes when predators arrived late and food was abundant. Fewer fish migrated downstream past a waterfall when food availability was high and in the presence of predators, and wild-genotype fry were more likely to migrate than fry of the fast-growing genotypes. After being returned to the experimental streams after migration, fast-growing genotypes survived equally well as those of the same genotypes that did not migrate, whereas migrating wild genotypes experienced higher mortality relative to those of the same genotypes that did not migrate. Comparisons of growth rates between siblings retained under hatchery conditions and those from habitats with the fastest growth in the experimental stream revealed that growth rates were similar for wild genotypes in both environments, whereas the fast-growing genotypes in the streams only realized 90% of their growth potential. The present study has shown that a major shift in developmental timing can alter critical early stages affecting survival and can have a significant effect on fitness. Furthermore, ecological conditions such as food abundance and predation pressure can strongly influence the potential for fast-growing variants to survive under natural conditions. The large-scale removal of many predatory species around the world may augment the evolution of increased intrinsic growth rates in some taxa.     

Experiments to test if allopatric Salvelinus alpinus are suitable year-round hosts of Gyrodactylus salaris (Monogenea)

There has been an increased focus on Salvelinus alpinus as a potential long-term host to Gyrodactylus salaris and, here, both susceptibility to G. salaris and ability to sustain a parasite population seasonally, was tested using fry and parr of S. alpinus from the River Skibotnelva, northern Norway. Fry were highly susceptible. Gyrodactylus salaris survived on allopatric S. alpinus parr during the 5 month-long winter when water temperatures were c. 1° C. Salvelinus alpinus fry also maintained a pulse of G. salaris infection for over 155 days from early May until autumn. Gyrodactylus salaris are thus able to reproduce and survive on S. alpinus for long periods and at low water temperatures. In spring, newly hatched fry of S. alpinus may serve as an important host to maintain a G. salaris metapopulation within a river system. The results suggest that S. alpinus are adequate long-term hosts of G. salaris independent of the presence of the co-occurring highly susceptible S. salar .     

Impact of herbivory and pollination on performance and competitive ability of oilseed rape transformed for pollen beetle resistance

Competitive ability of transgenic oilseed rape transformed with a pea lectin gene was estimated by comparisons of its performance when grown in a mixture with its non-transgenic counterpart and when grown singly, with and without pollen beetles present. The experiments were carried out in cages, once with bumblebees as pollinators and once without. In the absence of herbivory but with the presence of bumblebees, singly grown plant lines without lectin generally performed better than lines with lectin. Pollen beetles affected plant growth and reproduction, but there were no consistent differences between the lectin and non-lectin plant lines indicating that the transgenic trait did not protect plants from pest attack. Herbivory reduced the number of seeds when bumblebees were present. In the absence of bumblebees, however, plants produced more seeds with pollen beetles than without, indicating that some pollination was carried out by the beetles. Efficient pollination affected the competitive abilities of the lines; lectin lines were more competitive with bumblebees present and the reverse was true when bumblebees were absent. In the presence of bumblebees, lectin lines gained from being grown mixed with its non-transgenic counterpart. Because the transgenic plants expressed pea lectin in developing pollen it is possible that pollen quality in those plants was reduced, which may explain why the lectin lines had an advantage over non-lectin lines when exchange of pollen between the two plant types was facilitated by bumblebees.     

Growth, fecundity and competitive ability of transgenic Trifolium subterraneum subsp. subterraneum cv. Leura expressing a sunflower seed albumin gene

Ecological risk assessment is an important step in the production and commercialisation of transgenic plants. To date, however, most risk assessment studies have been performed on crop plants, and few have considered the ecological consequences associated with genetic modification of pasture species. In this study we compared the growth, yield, population dynamics and competitive ability of transgenic Trifolium subterraneum subsp. subterraneum cv. Leura (subclover) expressing a nutritive sunflower seed albumin (ssa) gene with the equivalent non-transgenic commercial line in a glasshouse competition trial. Plants were grown in low-fertility soil typical of unimproved native southeastern Australian grasslands. We measured survivorship, seed production rate, seed germination rate, seed weight, dry weight yield and the intrinsic rate of population increase (lambda) of plants grown in mixtures and monocultures over a range of densities (250 to 2000 plants m(-2)), and also determined intragenotypic and intergenotypic competition coefficients for each line. There were no significant differences between transgenic and non-transgenic plants in any of the measured variables except survivorship; transgenic plants had a significantly lower survival rate than non-transgenic plants when grown at high densities (p<0.01). However, density-dependent effects were observed for all measured variables, and in all models plant density affected the response variables more than the presence of the transgene. Based on these results, we conclude that the ssa gene construct appears to confer no advantage to transgenic T. s. subterraneum cv. Leura growing in mixed or pure swards under the fertility and density regimes examined in the trial. Our data also suggest that transgenic subterranean clover expressing the ssa gene is unlikely to exhibit a competitive advantage over associated non-transgenic commercial cultivars when grown in dense swards in low-fertility pastures.     

Aggression and growth depression in juvenile Atlantic salmon: the consequences of individual variation in standard metabolic rate

In aquaculture, competitive interactions for food are a major source of growth rate variation, since they result in aggressive individuals acquiring a disproportionate amount of food and growing faster. Consequently, such competition increases the variance and skew of the size distribution of fish. In Atlantic salmon Salmo salar acquaculture, rearing is often initiated with juveniles of uniform size. However, the initial factors allowing fish to out-compete others of the same size is unclear. This study shows that individual differences in standard metabolic rate (SMR) may contribute to differences in aggression between juvenile Atlantic salmon. Fish were segregated into three holding tanks on the basis of differences in relative SMR aggression was highest in the tank containing high SMR fish, and lowest in a low SMR group. However, there were no significant differences in mean growth between the three treatments, although the skew of the size distribution was much less in the low SMR group than in the high SMR group. Therefore, although mean growth was not improved in the low SMR group by excluding high SMR fish, their growth subsequently showed less variability, possibly as a consequence of fewer despotic individuals gaining disproportionate access to food.     

Food quality effects on instar‐specific life histories of a holometabolous insect

1. It is a long‐standing challenge to understand how changes in food resources impact consumer life history traits and, in turn, impact how organisms interact with their environment. To characterize food quality effects on life history, most studies follow organisms throughout their life cycle and quantify major life events, such as age at maturity or fecundity. From these studies, we know that food quality generally impacts body size, juvenile development, and life span. Importantly, throughout juvenile development, many organisms develop through several stages of growth that can have different interactions with their environment. For example, some parasitoids typically attack larger instars, whereas larval insect predators typically attack smaller instars. Interestingly, most studies lump all juvenile stages together, which ignores these ecological changes over juvenile development.
2. We combine a cross‐sectional experimental approach with a stage‐structured population model to estimate instar‐specific vital rates in the bean weevil, Callosobruchus maculatus across a food quality gradient. We characterize food quality effects on the bean weevil's life history traits throughout its juvenile ontogeny to test how food quality impacts instar‐specific vital rates.
3. Vital rates differed across food quality treatments within each instar; however, their effect differed with instar. Weevils consuming low‐quality food spent 38%, 37%, and 18% more time, and were 34%, 53%, and 63% smaller than weevils consuming high‐quality food in the second, third, and fourth instars, respectively. Overall, our results show that consuming poor food quality means slower growth, but that food quality effects on vital rates, growth and development are not equal across instars. Differences in life history traits over juvenile ontogeny in response to food quality may impact how organisms interact with their environment, including how susceptible they are to predation, parasitism, and their competitive ability.

     

Effects of infochemicals released by gape-limited fish on life history traits of Daphnia: a maladaptive response?

Life history shifts in daphnids in response to fish infochemicalsare generally interpreted as an adaptive response to positivesize-selective predation. This interpretation does, however,not hold for larval and small juvenile planktivorous fish, whichdue to gape limitation, feed on small and medium sized prey.In a life table experiment we show that daphnids exposed toinfochemicals excreted by small gape-limited perch and largerperch changed their life history in the same direction, irrespectiveof the contrasting size-selection of the fish. However, responsesto fish infochemicals were strongly influenced by food conditionsfor daphnids. In the high food treatments size at maturity wasin the presence of fish infochemicals, whereas age at maturityremained unchanged. Under low food conditions, size at maturitywas generally smaller compared with the high food situation,but unaffected by fish infochemicals. By contrast, age at maturity,which was increased at low food levels, was significantly lowerin fish treatments compared with the control. We conclude thatlife history responses of daphnids to gape-limited fish canindeed be maladaptive, but only in situations of high food availability.This combination of factors is, however, rather unlikely becausegape-limited fish usually occur in late spring during the clearwater phase when daphnids are severely food limited. We thushypothesize that the costs of this maladaptive response undernegative size-selective predation will be low under field conditionsand the selective advantage under positive size-selective predationlater in the season will outweigh these costs.     

Predator Avoidance of Transgenic Channel Catfish Containing Salmonid Growth Hormone Genes

Transgenic channel catfish (Ictalurus punctatus) containing salmonid growth hormone genes can grow 33% faster than normal channel catfish under aquaculture conditions. However, before transgenic catfish are released and utilized by the private sector, their genetic impact on the natural environment must be examined. Predator avoidance is one of the major fitness traits determining potential environmental risk. To determine the predator avoidance ability and growth performance of transgenic catfish in a natural habitat, various densities of transgenic and nontransgenic channel catfish were communally stocked in 0.04-ha earthen ponds without supplemental feeding. Largemouth bass (Micropterus salmoides) and green sunfish (Lepomis cyanellus) were stocked as predators. Nontransgenic fry had better predator avoidance than transgenic channel catfish when data were pooled (p < .01). When data were not pooled, nontransgenic catfish had better predator avoidance in six trials and transgenic individuals had better predator avoidance in four trials. There was no difference in predator avoidance in three trials. Overall predator avoidance was also better for nontransgenic individuals (p < .01) when the fish were evaluated as 3.5-g fingerlings, more clearly than as fry, as transgenic individuals were more vulnerable in 3 of 4 trials at this life stage. There was no significant difference in growth performance between transgenic and nontransgenic channel catfish in ponds without supplemental feeding. These findings indicate that transgenic channel catfish could be used for commercial aquaculture without affecting the natural environment. Although transgenic channel catfish may be released to nature by accident, any ecological effect would be unlikely because the increased susceptibility of transgenic channel catfish to predators would most likely decrease or eliminate the transgenic genotype. Received March 8, 1999; accepted June 3, 1999.     

抗真菌转基因水稻秸秆降解对土壤细菌数量及多样性的影响

【背景】随着转基因作物的大面积种植,其潜在的环境安全性问题备受关注。转基因作物收获后,大部分残留物会重返土壤,可能对土壤微生物造成影响。【方法】通过室内模拟田间秸秆降解试验,采用平板计数法、表面荧光显微镜直接计数法（FDC）以及变性梯度凝胶电泳（DGGE）技术,分析了抗真菌转基因水稻秸秆降解对土壤细菌数量及多样性的影响。【结果】平板计数表明,在整个降解过程中,转基因与非转基因处理土壤可培养细菌数量的变化趋势有所差别,但差异不显著。FDC结果表明,转基因与非转基因处理土壤细菌总数差异不显著。DGGE指纹图谱显示,转基因与非转基因处理土壤样品之间的多样性指数、均匀度和丰富度均无显著差异。【结论与意义】抗真菌转基因水稻秸秆降解并未对土壤细菌数量和多样性产生显著影响。本研究为抗真菌转基因水稻的环境安全性评估提供了依据。     

Costs of compensation: effect of early life conditions and reproduction on flight performance in zebra finches

Conditions experienced in early life have been shown to affect the development or programming of physiological processes. While animals may recover from earlier periods of adversity, this process can carry long-term costs. Such long-term effects are likely to be most evident when individuals are placed in demanding situations that require high performance. Escape flight speed in passerine birds is crucial to predator evasion and requires very rapid take-off. Here, we examine whether the ability to maintain escape flight performance during the immediate post-breeding period is influenced by conditions in early life. We manipulated the early life conditions experienced by zebra finches (Taeniopygia guttata) by rearing them on either low or high quality food through the growth period, or by changing conditions halfway through the nestling period, moving from high to low or vice versa. While there was no difference amongst the treatment groups in body size attained by adulthood, amongst the birds that experienced low quality food, the body size of those that were switched to a high quality diet halfway through the nestling growth period recovered faster than those that had low quality food until fledging. We found no differences amongst the dietary groups in flight performance at adulthood prior to breeding, and all groups showed a decline in average escape flight performance over the breeding period. However, the magnitude of the post-breeding decline in flight performance for a given level of reproductive output was significantly greater for those females that had experienced a switch from a low to a high quality diet during the nestling phase. These results suggest that this diet-induced rapid recovery of body size, which may have immediate competitive advantages, nonetheless carries locomotory costs in later life manifest in the capacity to sustain the high performance escape response during the post-reproductive recovery phase.     

The evolution of growth rate, resource allocation and competitive ability in seeder and resprouter tree seedlings

Many woody plant species in fire disturbed communities survive disturbance events by resprouting. The resprouting life history is predicted to be costly to plants as resources are diverted into storage for post-fire regrowth rather than allocated to current growth, and resprouting species typically grow more slowly than seeder species (species that do not resprout after disturbance events). Differences in allocation to current growth are also predicted to make resprouter species poorer competitors compared to seeder species. We tested the predictions that the evolution of a resprouter life history is associated with slow growth, increased allocation to storage, and low competitive ability in woody plant seedlings. We grew eight phylogenetically independent pairs of seeder and resprouter species in competition and no competition treatments in a field experiment near Sydney, Australia. The presence of competitors reduced plant growth rates across taxa and fire response life histories. However, relative to seeder species, resprouter species were not slower growing, they did not allocate more resources to storage, and they did not have lower competitive abilities. We propose that differences in resource allocation to storage are not responsible for differences in growth rate and competitive ability. Rather, growth rate and competitive ability in seedlings are associated with key aspects of plant life history such as life-span and body size at maturity. These traits that are sometimes, but not always, related to fire response life histories.     

Interactions between riparian shading and food supply: a seasonal comparison of effects on time budgets, space use and growth in Atlantic salmon Salmo salar

This study examines seasonal (winter v. summer) differences in space-time budgets, food intake and growth of Atlantic salmon Salmo salar parr in a controlled, large-scale stream environment, to examine the direction and magnitude of shifts in behaviour patterns as influenced by the availability of overhead cover and food supply. Salmo salar parr tested in the presence of overhead cover were significantly more nocturnal and occupied more peripheral positions than those tested in the absence of overhead cover. This increase in nocturnal activity was driven primarily by increased activity at night, accompanied by a reduction in daytime activity during winter. The presence of overhead cover had no effect on rates of food intake or growth for a given food supply in a given season. Growth rates were significantly higher for fish subjected to a high food supply than those subjected to a low food supply. Food supply did not affect the extent to which S. salar parr were nocturnal. These results were consistent between winter and summer. The use of riparian shading as a management technique to mitigate the effects of warming allows the adoption of more risk-averse foraging behaviour and may be particularly beneficial in circumstances where it serves also to increase the availability of food.     

Gall insects and selection on plant vigor: can susceptibility compromise success in competition?

Gall insects select vigorously growing plants and plant parts when initiating gall formation. Vigor is associated with rapid growth rate, and in turn, rapid growth confers competitiveness. Are there conditions under which the cost of vigor, in the form of increased susceptibility to attack, outweighs the benefit of competitive success? I present a simulation model to explore the interaction between susceptibility and competition on the selective advantage of increased growth rate. Assuming size-symmetric competition, the model shows that in general, vigor is favored (benefit > cost) at low to intermediate gall loads. At very high plant densities, however, plants with high gall loads may lose standing in the competitive size hierarchy from which they cannot recover. The details of this result, however, change somewhat when competition is size-asymmetric, that is, when a larger focal plant suppresses smaller neighbors, but smaller neighbors cannot exert a reciprocal effect on the focal. At low densities, the pattern of selection on growth rate is qualitatively similar to the size-symmetric case. At higher plant densities, however, fast-growing genotypes can suppress slow ones so much during the preattack phase that even at the highest gall loads they maintain their standing in the competitive hierarchy. Thus, heavy gall insect attack on vigorous plants can impose selection against high intrinsic growth rates under strong symmetric competition, but not strong asymmetric competition. While life history traits can evolve as a correlated response to selection on defensive traits that reduce susceptibility, this model reveals that susceptibility can evolve as a correlated response to selection on basic life history traits.