Species recognition is an important aspect of an organism's biology. Here, we consider how parasitoid wasps vary their reproductive decisions when their offspring face intra- and interspecific competition for resources and mates. We use host acceptance and sex ratio behaviour to test whether female Nasonia vitripennis and Nasonia longicornis discriminate between conspecifics and heterospecifics when ovipositing. We tested pairs of conspecific or heterospecific females ovipositing either simultaneously or sequentially on a single host, using strains varying in their recent history of sympatry. Both N. vitripennis and N. longicornis rejected parasitized hosts more often than unparasitized hosts, although females were more likely to superparasitize their own species in the sequential treatment. However, sex ratio behaviour did not vary, suggesting similar responses towards conspecifics and heterospecifics. This contrasts with theory predicting that heterospecifics should not influence sex ratios as their offspring do not influence local mate competition, where conspecifics would. These non-adaptive sex ratios reinforce the lack of adaptive kin discrimination in N. vitripennis and suggest a behavioural constraint. Discrimination between closely related species is therefore context dependent in Nasonia. We suggest that isolating mechanisms associated with the speciation process have influenced behaviour to a greater extent than selection on sex ratios.
species recognition; speciation; adaptation; sex ratios; superparasitism; multiparasitism
The acquisition and use of information are essential for decision-making in an uncertain world. The use of social information, or information from the behaviour of others, may be a common and efficient mechanism to improve estimates of resource quality by animals. According to theory, social information cues with higher information content should have a greater influence on decision-making, and current information should be weighed more than prior information. However, experimental tests of these hypotheses remain scarce. We exposed female cactus bugs (Chelinidea vittiger) to different types of social information (the presence of conspecific eggs or nymphs) presented at different times (current or prior to egg laying) to determine the influence of social information on offspring production. We found that social information substantially altered the number of eggs produced. The presence of conspecific eggs, regardless of timing, consistently increased egg production, whereas nymphs only increased egg production when present during egg laying. We conclude that the type and timing of social information may have an important, yet unappreciated, influence on reproductive allocation.
prior information; public information; reproductive investment; oviposition
Many species disperse during their lifetime. Two factors that can affect the performance of individuals following dispersal are the presence of conspecifics and intrinsic habitat quality at the settlement site. Detecting the influence of these factors can be difficult for at least two reasons: (1) the outcomes of interactions with conspecifics are often variable including both competition and facilitation, and (2) selection of high quality habitats often leads to positive covariance between habitat quality and density. In this study, I investigate positive and negative effects of resident blue streak cleaner wrasse (Labroides dimidiatus) on the growth and survival of recently settled conspecifics while accounting for habitat quality. Juvenile L. dimidiatus settle near adult conspecifics, but likely have to compete with resident adults for access to food. However, field experiments indicate that settlers have access to more resources at occupied sites, and as a result, grow faster despite evidence for competition with residents. This result is a direct consequence of two factors: (1) resident conspecifics facilitate settlers by attracting client fish, and (2) resident conspecifics are strongly associated with high quality habitat. These results highlight the need to simultaneously consider habitat quality and competitive and facilitative interactions between conspecifics when making inferences about ecological processes from spatial patterns of individual performance.
Electronic supplementary material
The online version of this article (doi:10.1007/s00442-010-1826-7) contains supplementary material, which is available to authorized users.
Density-dependent growth; Habitat selection; Labroides dimidiatus; Coral reef; Fish
Many insects deposit marking pheromones following egg-laying that signal an occupied and thus sub-optimal resource. Herbivorous insects mark host fruit or other vegetative plant parts after depositing eggs, while insect parasitoids deposit such pheromones directly on the cuticle of a particular life stage of their prey. These oviposition marking pheromones (OMPs) are then recognized by conspecifics, which avoid subsequent egg-laying in the previously utilized and unsuitable host. Since many host resources are capable of supporting a limited number of offspring, these pheromones function to decrease competition among the brood, which increases survival rate of the subsequent generation. In rare instances, distinct species of phytophagous and parasitic insects will inspect the same substrate following egg-laying.1 Recently, Stelinski et al.1 have demonstrated that in such instances, the herbivore is able to learn to recognize its predator's OMP and utilize it to its advantage by avoiding oviposition into unsuitable host fruit. This recognition of a foreign marking pheromone occurs in a multitrophic context since both herbivore and parasitoid inspect, oviposit into, and mark the same substrate (i.e., fruit surface). In this Article Addendum, we further show that this recognition of a foreign pheromone is both context-dependent and mediated by preimaginal conditioning.
xenodeictic pheromone; Rhagoletis pomonella; Diachasma alloeum; host marking; parasitoid; behavioral plasticity; larval conditioning
Females of many species can gain benefits from being choosy about their mates and even exhibit context-dependent investment in reproduction in response to the quality of their breeding situation. Here, we show that if a male house wren is provided with surplus nest boxes in his territory, his mate lays a larger clutch with a significantly higher proportion of sons. This response to a territory characteristic directly associated with male competitive ability, and ultimately to male reproductive success, suggests that male competition over access to high-quality territories with surplus nest boxes (i.e. those able to support polygyny) may influence female reproductive investment decisions. The results of this study have interesting implications, particularly considering the important role that studies of cavity nesting birds utilizing nest boxes have played in advancing our understanding of behaviour, ecology and evolution.
clutch size; differential allocation; offspring sex ratio; parental investment; sexual selection; Troglodytes aedon
Conspecifics are usually considered competitors negatively affecting food intake rates. However, their presence can also inform about resource quality by providing inadvertent social information. Few studies have investigated whether foragers perceive conspecifics as informers or competitors. Here, we experimentally tested whether variation in the density of demonstrators (‘none’, ‘low’ and ‘high’), whose location indicated flower profitability, affected decision-making of bumble-bees Bombus terrestris. Bumble-bees foraged on either ‘simple’ (two colours) or ‘complex’ (four colours) artificial floral communities. We found that conspecifics at low density may be used as sources of information in first flower choices, whereas they appeared as competitors over the whole foraging sequence. Low conspecific densities improved foragers' first-visit success rate in the simple environment, and decreased time to first landing, especially in the complex environment. High conspecific densities did not affect these behavioural parameters, but reduced flower constancy in both floral communities, which may alter the efficiency of pollinating visits. These results suggest that the balance of the costs and benefits of conspecific presence varies with foraging experience, floral community and density. Spatio-temporal scales could thus be an important determinant of social information use. This behavioural flexibility should allow bumble-bees to better exploit their environment.
Bombus terrestris; decision-making; competition; foraging behaviour; inadvertent social information; plant–pollinator interactions
This study investigated the egg-laying behaviour of ectoparsitoid, Dinarmus basalis Rondani (Hymenoptera: Pteromalidae), females when faced with a prolonged deprivation of suitable hosts leading to extreme ‘oviposition pressure’. The egg-laying behaviour of virgin D. basalis females was tested with Callosobruchus maculatus (F.) (Coleoptera: Bruchidae) hosts previously parasitized by the conspecific females in which the developing larvae had reached the last larval instar (L5) or pupae. The hyperparasitism did not prevent the occurrence of superparasitism, but only one D. basalis egg from a hyperparasitized D. basalis L5 larvae reached the adult stage due to the solitary behaviour of the D. basalis larvae. Under these experimental conditions, 60.78% of the D. basalis adults emerging from larvae were miniaturized due to the depletion of host resources.
Callosobruchus maculatus; Eupelmus vuilleti; Eupelmus orientalis; Vigna unguiculata; Bruchidae; intraspecific; interspecific discrimination; intraspecific competition; cleptoparasitic behaviour
Effects of diet on longevity are complex because acquired resources are shared among growth, reproduction and somatic maintenance. We simplify these axes by examining how dietary restriction and competitive contexts affect longevity using semelparous males of the Australian redback spider (Latrodectus hasselti). Plastic development of L. hasselti males results in trade-offs of body condition against faster development if females are present, facilitating scramble competition. In the absence of females, males develop slowly as high body condition adults, and are better equipped for mate searching. Here we focus on effects of diet and competitive context on body condition and longevity. Although male survival depended on body condition and exercise, contrary to studies in a wide range of taxa, dietary restriction did not increase longevity. However, there was an interactive effect of diet and competitive context on lifespan, because high-diet males reared in the absence of females lived longer than males reared in the presence of females. Thus males near females pay a survival cost of developing rapidly. This shows that life-history trade-offs affected by competitive context can impose longevity costs independent of the direct energy expenditure of searching, courtship, competition or reproduction.
dietary restriction; life-history trade-offs; longevity; developmental plasticity; Latrodectus hasselti
Animals are active at different times of the day and their activity schedules are shaped by competition, time-limited food resources and predators. Different temporal niches provide different light conditions, which affect the quality of visual information available to animals, in particular for navigation. We analysed caste-specific differences in compound eyes and ocelli in four congeneric sympatric species of Myrmecia ants, with emphasis on within-species adaptive flexibility and daily activity rhythms. Each caste has its own lifestyle: workers are exclusively pedestrian; alate females lead a brief life on the wing before becoming pedestrian; alate males lead a life exclusively on the wing. While workers of the four species range from diurnal, diurnal-crepuscular, crepuscular-nocturnal to nocturnal, the activity times of conspecific alates do not match in all cases. Even within a single species, we found eye area, facet numbers, facet sizes, rhabdom diameters and ocelli size to be tuned to the distinct temporal niche each caste occupies. We discuss these visual adaptations in relation to ambient light levels, visual tasks and mode of locomotion.
temporal niches; compound eyes; ocelli; castes; workers; alates
Understanding how individuals select mates becomes complex when high-quality conspecifics resemble heterospecifics. Individuals facing such a situation may be unable to effectively identify both conspecifics (species recognition) and high-quality mates that can confer fitness benefits to the choosy individual or its offspring (mate-quality recognition). Here I suggest when a conflict may occur between species and mate-quality recognition, discuss the evolutionary consequences stemming from this conflict, and present a model of mate-preference evolution in response to heterospecifics. Determining how species and mate-quality recognition interact to shape mate-choice decisions is important for understanding the diversification of sexually selected traits among closely related taxonomic groups, the use of complex sensory systems for detecting mates, and seemingly inappropriate mate-choice decisions.
Sperm competition occurs when 2 or more males copulate with a particular female during the same reproductive cycle, and their sperm compete to fertilize the female's available eggs. One strategy that male voles use to assess the risk and intensity of sperm competition involves responding to the presence of scent marks of conspecific males found near a sexually receptive female. Previously, we have shown that if a male vole copulated with a female while he was in the presence of the odors of another male he increased his sperm investment relative to his investment if another male's odors were not present. The aim of the present study was to test the hypothesis that males assess differences in the relative quality of competing males and adjust their sperm investment accordingly. We did so by allowing males to copulate when they were exposed to the scent mark of a 24-h food-deprived male (low-quality male) or the scent mark of a male that was not food deprived (high-quality male). The data indicate that male meadow voles did not increase their sperm investment during copulation when exposed to the scent mark of a food-deprived male but did so when they were exposed to the scent mark of a male that was not food deprived. The results support the hypothesis that male voles are able to adjust sperm investment when they encounter the scent marks of males that differ in quality.
chemical signals; copulatory behavior; food deprivation; scent marking; sperm competition; voles
Conspecific competition occurs in a multitude of organisms, particularly in parasites, where several clones are commonly sharing limited resources inside their host. In theory, increased or decreased transmission investment might maximize parasite fitness in the face of competition, but, to our knowledge, this has not been tested experimentally. We developed and used a clone-specific, stage-specific, quantitative PCR protocol to quantify Plasmodium chabaudi replication and transmission stage densities in mixed-clone infections. We co-infected mice from two strains with an avirulent and virulent parasite clone and found competitive suppression of in-host (blood-stage) parasite densities and generally corresponding reductions in transmission stage production, with the virulent clone obtaining overall competitive superiority. In response to competitive suppression, there was little evidence of any alteration in transmission stage investment, apart from a small reduction by one of the two clones in one of the two host strains. This alteration did not result in a competitive advantage, although it might have reduced the disadvantage. This study supports much of the current literature, which predicts that conspecific in-host competition will result in a competitive advantage and positive selection for virulent clones and thus the evolution of higher virulence.
virulence; quantitative RT-PCR; life history; transmission; gametocyte; phenotypic plasticity
Rather than being a static, species specific trait, reproductive behavior in female amphibians is variable within an individual during the breeding season when females are capable of reproductive activity. Changes in receptivity coincide with changes in circulating estrogen. Estrogen is highest at the point when females are ready to choose a male and lay eggs. At this time female receptivity (her probability of responding to a male vocal signal) is highest and her selectivity among conspecific calls (measured by her probability of responding to a degraded or otherwise usually unattractive male signal) is lowest. These changes occur even though females retain the ability to discriminate different acoustic characteristics of various conspecific calls. After releasing her eggs, female amphibians quickly become less receptive and more choosy in terms of their responses to male sexual advertisement signals. Male vocal signals stimulate both behavior and estrogen changes in amphibian females making mating more probable. The changes in female reproductive behavior are the same as those generally accepted as indicative of a change in female sexual arousal leading to copulation. They are situationally triggered, gated by interactions with males, and decline with the consummation of sexual reproduction with a chosen male. The changes can be triggered by either internal physiological state or by the presence of stimuli presented by males, and the same stimuli change both behavior and physiological (endocrine) state in such a way as to make acceptance of a male more likely. Thus amphibian females demonstrate many of the same general characteristics of changing female sexual state that in mammals indicate sexual arousal.
amphibians; reproduction; female; communication; estrogen
Among the extraordinary adaptations driven by sperm competition is the cooperative behaviour of spermatozoa1. By forming cooperative groups, sperm can increase their swimming velocity and thereby gain an advantage in intermale sperm competition1,2. Accordingly, selection should favour cooperation of the most closely related sperm to maximize fitness3. Here we show that sperm of deer mice (genus Peromyscus) form motile aggregations, then we use this system test predictions of sperm cooperation. We first show that sperm aggregate more often with conspecific than heterospecific sperm, suggesting that individual sperm can discriminate based on genetic relatedness. Next, we provide evidence that the cooperative behaviour of closely-related sperm is driven by sperm competition. In a monogamous species lacking sperm competition, P. polionotus, sperm indiscriminately group with unrelated conspecific sperm. In contrast, in the highly promiscuous deer mouse, P. maniculatus, sperm are significantly more likely to aggregate with those obtained from the same male than sperm from an unrelated conspecific donor. Even when we test sperm from sibling males, we continue to see preferential aggregations of related sperm in P. maniculatus. These results suggest that sperm from promiscuous deer mice discriminate among relatives and thereby cooperate with the most closely-related sperm, an adaptation likely driven by sperm competition.
Genomic imprinting has been proposed to evolve when a gene's expression has fitness consequences for individuals with different coefficients of matrilineal and patrilineal relatedness, especially in the context of competition between offspring for maternal resources. Previous models have focused on pre-emptive hierarchies, where conflict arises with respect to resource allocation between present and future offspring. Here we present a model in which imprinting arises from scramble competition within litters. The model predicts paternal-specific expression of a gene that increases an offspring's fractional share of resources but reduces the size of the resource pool, and maternal-specific expression of a gene with opposite effects. These predictions parallel the observation in economic models that individuals tend to underprovide public goods, and that the magnitude of this shortfall increases with the number of individuals in the group. Maternally derived alleles are more willing than their paternally derived counterparts to contribute to public goods because they have a smaller effective group size.
The use of ‘tactical deception’ is argued to have been important in the cognitive evolution of the order Primates, but systematic studies of active deception in wild non-human primates are scant. This study tests whether wild tufted capuchin monkeys (Cebus apella nigritus) use alarm calls in a functionally deceptive manner to usurp food resources. If capuchins use alarm calls ‘deceptively’, it was predicted that false alarms should be: (i) given by subordinates more than by dominants, (ii) more frequent when food is most contestable, (iii) more frequent when less food is available, and (iv) given when the caller is in a spatial position in which it could increase its feeding success if conspecifics react to the call. These predictions were tested by observing subjects in experimental contexts, in which the amount and distribution of a high-value resource (banana pieces) were manipulated using wooden platforms suspended from tree branches. While false alarms were non-significantly more common when more food was available, the three remaining predictions were supported. These results generally support the hypothesis that alarm calls are used by capuchins to reduce the effects of feeding competition. Whether this is intentional on the part of the caller requires further investigation.
alarm calls; feeding competition; deception; communication; primates; Cebus apella nigritus
Plasticity in reproductive physiology is one avenue by which environmental signals, such as poor quality food, can be coordinated with adaptive responses. Insects have the ability to resorb oocytes that are not oviposited. Oosorption is proposed to be an adaptive mechanism to optimize fitness in hostile environments, recouping resources that might otherwise be lost, and reinvesting them into future reproductive potential. We tested the hypothesis that oosorption is an evolved mechanism by which females can reallocate resources from current reproductive effort to survival and future reproduction, when conditions for reproduction are poor, by examining the reproductive physiology and life-history outcome under poor quality food in populations of the milkweed bug (Oncopeltus fasciatus) that have adapted to live on sunflower seed. Females fed a diet of pumpkin seeds, known to be a poor host food, had higher levels of ovarian apoptosis (oosorption), lower reproductive output, but no reduction in life span under poor nutrition, as predicted under the oosorption hypothesis. However, the schedule of reproduction was surprising given the “wait to reproduce” assumption of oosorption as early fecundity was unaffected.
Energy allocation; nutrition; oosorption; ovarian apoptosis; reproductive investment
Resource competition is thought to play a major role in driving evolutionary diversification. For instance, in ecological character displacement, coexisting species evolve to use different resources, reducing the effects of interspecific competition. It is thought that a similar diversifying effect might occur in response to competition among members of a single species. Individuals may mitigate the effects of intraspecific competition by switching to use alternative resources not used by conspecific competitors. This diversification is the driving force in some models of sympatric speciation, but has not been demonstrated in natural populations. Here, we present experimental evidence confirming that competition drives ecological diversification within natural populations. We manipulated population density of three-spine sticklebacks (Gasterosteus aculeatus) in enclosures in a natural lake. Increased population density led to reduced prey availability, causing individuals to add alternative prey types to their diet. Since phenotypically different individuals added different alternative prey, diet variation among individuals increased relative to low-density control enclosures. Competition also increased the diet–morphology correlations, so that the frequency-dependent interactions were stronger in high competition. These results not only confirm that resource competition promotes niche variation within populations, but also show that this increased diversity can arise via behavioural plasticity alone, without the evolutionary changes commonly assumed by theory.
diversification; Gasterosteus aculeatus; individual specialization; intraspecific competition; optimal foraging theory; negative frequency dependence
Understanding the size of clutches produced by only one parent may require a game-theoretic approach: clutch size may affect offspring fitness in terms of future competitive ability. If larger clutches generate smaller offspring and larger adults are more successful in acquiring and retaining resources, clutch size optima should be reduced when the probability of future competitive encounters is higher. We test this using Goniozus nephantidis, a gregarious parasitoid wasp in which the assumption of size-dependent resource acquisition is met via female–female contests for hosts. As predicted, smaller clutches are produced by mothers experiencing competition, due to fewer eggs being matured and to a reduced proportion of matured eggs being laid. As assumed, smaller clutches generate fewer but larger offspring. We believe this is the first direct evidence for pre-ovipositional and game-theoretic clutch size adjustment in response to an intergenerational fitness effect when clutches are produced by a single individual.
clutch size; game-theory; contests; intergenerational fitness
Conspecific sperm precedence (CSP) has been identified as an important post-copulatory, pre-zygotic mechanism that can act to reduce gene flow between populations. The evolution of CSP is thought to have arisen as a by-product of male and female coevolution in response to intraspecific post-copulatory sexual selection. However, little is known about the mechanisms that generate CSP. When Callosobruchus subinnotatus females copulate with both C. subinnotatus and Callosobruchus maculatus males, regardless of mating order, the majority of eggs are fertilized by conspecific sperm. The low number of heterospecific fertilizations does not result from general differences in the viability of sperm in the female reproductive tract, as heterospecific sperm fertilized equivalent numbers of eggs as conspecific sperm in the absence of sperm competition. Instead, CSP results from disadvantages to heterospecific sperm that are manifest only when in competition with conspecific sperm. CSP in C. subinnotatus appears to result from two, not mutually exclusive, mechanisms. First, conspecific sperm are better able to displace heterospecific sperm from female storage. Second, conspecific sperm achieve disproportionately higher numbers of fertilizations relative to their proportional representation in the fertilization set. Thus, we provide evidence of differential sperm use from the female spermatheca.
conspecific sperm precedence; post-copulatory sexual selection; sperm selection; Callosobruchus
Evolutionary theory predicts that female intrasexual competition will occur when males of high genetic quality are considered to be a resource. It is probable that women compete in terms of attractiveness since this is one of the primary criteria used by men when selecting mates. Furthermore, because hormones influence the mate-selection process, they may also mediate competition. One competitive strategy that women use is derogation--any act intended to decrease a rival's perceived value. To investigate intrasexual competition through derogation, the influence of oestrogen on women's ratings of female facial attractiveness was examined. During periods of high oestrogen, competition, and hence derogation, increased, as evidenced by lower ratings of female facial attractiveness. By contrast, oestrogen levels did not significantly affect ratings of male faces. These findings support the theory of female intrasexual competition with respect to attractiveness.
When selecting a resource to exploit, an insect colony must take into account at least two constraints: the resource must be abundant enough to sustain the whole group, but not too large to limit exploitation costs, and risks of conflicts with other colonies. Following recent results on cockroaches and ants, we introduce here a behavioral mechanism that satisfies these two constraints. Individuals simply modulate their probability to switch to another resource as a function of the local density of conspecifics locally detected. As a result, the individuals gather at the smallest resource that can host the whole group, hence reducing competition and exploitation costs while fulfilling the overall group's needs. Our analysis reveals that the group becomes better at discriminating between similar resources as it grows in size. Also, the discrimination mechanism is flexible and the group readily switches to a better suited resource as it appears in the environment. The collective decision emerges through the self-organization of individuals, that is, in absence of any centralized control. It also requires a minimal individual cognitive investment, making the proposed mechanism likely to occur in other social species and suitable for the development of distributed decision making tools.
Sperm competition theory predicts that males should increase their expenditure on the ejaculate with increasing risk of sperm competition, but decrease their expenditure with increasing intensity. There is accumulating evidence for sperm competition theory, based on examinations of testes size and/or the numbers of sperm ejaculated. However, recent studies suggest that ejaculate quality can also be subject to selection by sperm competition. We used experimental manipulations of the risk and intensity of sperm competition in the cricket, Teleogryllus oceanicus. We found that males produced ejaculates with a greater percentage of live sperm when they had encountered a rival male prior to mating. However, when mating with a female that presented a high intensity of sperm competition, males did not respond to risk, but produced ejaculates with a reduced percentage of live sperm. Our data suggest that males exhibit a fine-tuned hierarchy of responses to these cues of sperm competition.
strategic ejaculation; ejaculate quality; sperm viability
Adult individuals of many species kill unrelated conspecific infants for several adaptive reasons ranging from predation or resource competition to the prevention of misdirected parental care. Moreover, infanticide can increase the reproductive success of the aggressor by killing the offspring of competitors and thereafter mating with the victimized females. This sexually selected infanticide predominantly occurs in polygynous species, with convincing evidence for primates, carnivores, equids, and rodents. Evidence for bats was predicted but lacking.
Here we report the first case, to our knowledge, of sexually selected infanticide in a bat, the polygynous white-throated round-eared bat, Lophostoma silvicolum. Behavioral studies in a free-living population revealed that an adult male repeatedly attacked and injured the pups of two females belonging to his harem, ultimately causing the death of one pup. The infanticidal male subsequently mated with the mother of the victimized pup and this copulation occurred earlier than any other in his harem.
Our findings indicate that sexually selected infanticide is more widespread than previously thought, adding bats as a new taxon performing this strategy. Future work on other bats, especially polygynous species in the tropics, has great potential to investigate the selective pressures influencing the evolution of sexually selected infanticide and to study how infanticide impacts reproductive strategies and social structures of different species.
Plants are sessile and therefore have to perceive and adjust to changes in their environment. The presence of neighbours leads to a competitive situation where resources and space will be limited. Complex adaptive responses to such situation are poorly understood at the molecular level.
Using microarrays, we analysed whole-genome expression changes in Arabidopsis thaliana plants subjected to intraspecific competition. The leaf and root transcriptome was strongly altered by competition. Differentially expressed genes were enriched in genes involved in nutrient deficiency (mainly N, P, K), perception of light quality, and responses to abiotic and biotic stresses. Interestingly, performance of the generalist insect Spodoptera littoralis on densely grown plants was significantly reduced, suggesting that plants under competition display enhanced resistance to herbivory.
This study provides a comprehensive list of genes whose expression is affected by intraspecific competition in Arabidopsis. The outcome is a unique response that involves genes related to light, nutrient deficiency, abiotic stress, and defence responses.
Arabidopsis thaliana; Gene expression; Intraspecific competition; Herbivory; Abiotic stress; Biotic stress