Extrinsic Competition Research Articles

Extrinsic and Intrinsic Competition between Chouioa cunea Yang and Tetrastichus septentrionalis (Hymenoptera: Eulophidae), Two Pupal Parasitoids of the Fall Webworm, Hyphantria cunea (Lepidoptera: Erebidae).

The endoparasitoids Chouioa cunea Yang and Tetrastichus septentrionalis Yang (Hymenoptera: Eulophidae) are both gregarious pupal parasitoids of the fall webworm, Hyphantria cunea (Drury) (Lepidoptera: Erebidae). In order to analyze the competitive interactions between both parasitoids exploiting H. cunea, we assessed both extrinsic and intrinsic competition. The search time, oviposition duration, and oviposition frequency were used as evaluation criteria for extrinsic competition. The number of survival days, female ratio, and number of parasitoids emerging from the host were used as evaluation criteria for intrinsic competition. The results indicated that both parasitoid species were able to parasitize hosts that were already parasitized by competitors. The first released species consistently emerged as the superior competitor in multiparasitized hosts. Both parasitoid release orders and time intervals between oviposition affect the competition of parasitoids and the parasitic efficiency. The results emphasize the parasitic abilities of both parasitoid species and provide a basis for future research on competition mechanisms and biological control of H. cunea.

Interspecific competition between Trissolcus japonicus and Trissolcus mitsukurii, two promising candidates for biocontrol of Halyomorpha halys

The invasive stink bug Halyomorpha halys, native to east Asia, is a severe agricultural pest of worldwide importance and biocontrol is a promising method for its long-term management. The two Asian egg parasitoids Trissolcus japonicus and Trissolcus mitsukurii are considered to be effective candidates as biological control agents. These species can co-occur in the same habitats and interspecific competition for hosts could therefore have an important role in natural control. We assessed the foraging behavior and the progeny production of both species under laboratory conditions with three different interspecific competition scenarios. With indirect competition, H. halys egg masses previously parasitized by the female of one species were offered to the competitor females after 0, 3, 5, or 7 days. With direct extrinsic competition, females of each species were tested in arenas either when introduced simultaneously (synchronous release) or when half of the host egg mass had already been parasitized by the other species (asynchronous release). Both Trissolcus species were able to parasitize host eggs already parasitized by the other species, although the progeny production always favored the species that arrived first on the host egg mass. However, in the synchronous releases scenario, T. mitsukurii displayed more aggressive behavior and spent more time defending the host egg mass than T. japonicus, resulting in a higher progeny production. Our results showed that multiparasitism among T. japonicus and T. mitsukurii, at least under laboratory conditions, resulted in a general waste of searching time, energy, eggs laid, and a lower reproductive potential by the second-arriving species on the host egg mass.

Contrasting reproductive traits of competing parasitoids facilitate coexistence on a shared host pest in a biological control perspective

BACKGROUNDInterspecific competition in insect parasitoids is an important ecological phenomenon that has relevant implications for biological pest control. To date, interspecific intrinsic (=larval) competition has been intensively studied, while investigations on extrinsic (=adult) competition have often lagged behind. In this study we examined the role played by parasitoid reproductive traits and host clutch size on the outcome of extrinsic competition between Trissolcus basalis (Wollaston) and Ooencyrtus telenomicida (Vassiliev), two egg parasitoids of the pest Nezara viridula (L). Laboratory experiments were conducted by allowing both parasitoid species to exploit an egg mass made of 10, 20, 30, or 40 hosts through single or simultaneous releases. Furthermore, under field conditions, egg masses consisting of 10 or 40 hosts were exposed in a tomato crop in order to validate laboratory investigation.RESULTSThe results show that the egg mass size is an important predictor of extrinsic competition in our study system as a higher proportion of T. basalis emerged from large egg masses, while O. telenomicida dominated in small egg masses. Analysis of reproductive traits of parasitoid species indicates that T. basalis has superior abilities in host exploitation compared with O. telenomicida.CONCLUSIONSWe found that contrasting reproductive traits of two competing egg parasitoid species facilitate coexistence on a shared stink bug host. This work also highlights the importance to consider extrinsic competitive interactions between parasitoid species in a biological control perspective. © 2022 The Authors. Pest Management Science published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.

Gamification suffers from the novelty effect but benefits from the familiarization effect: Findings from a longitudinal study

There are many claims that gamification (i.e., using game elements outside games) impact decreases over time (i.e., the novelty effect). Most studies analyzing this effect focused on extrinsic game elements, while fictional and collaborative competition have been recently recommended. Additionally, to the best of our knowledge, no long-term research has been carried out with STEM learners from introductory programming courses (CS1), a context that demands encouraging practice and mitigating motivation throughout the semester. Therefore, the main goal of this work is to better understand how the impact of a gamification design, featuring fictional and competitive-collaborative elements, changes over a 14-week period of time, when applied to CS1 courses taken by STEM students (N = 756). In an ecological setting, we followed a 2x7 quasi-experimental design, where Brazilian STEM students completed assignments in either a gamified or non-gamified version of the same system, which provided the measures (number of attempts, usage time, and system access) to assess user behavior at seven points in time. Results indicate changes in gamification’s impact that appear to follow a U-shaped pattern. Supporting the novelty effect, the gamification’s effect started to decrease after four weeks, decrease that lasted between two to six weeks. Interestingly, the gamification’s impact shifted to an uptrend between six and 10 weeks after the start of the intervention, partially recovering its contribution naturally. Thus, we found empirical evidence supporting that gamification likely suffers from the novelty effect, but also benefits from the familiarization effect, which contributes to an overall positive impact on students. These findings may provide some guidelines to inform practitioners about how long the initial contributions of gamification last, and how long they take to recover after some reduction in benefits. It can also help researchers to realize when to apply/evaluate interventions that use gamification by taking into consideration the novelty effect and, thereby, better understand the real impact of gamification on students’ behavior in the long run.

Intrinsic and Extrinsic Competition: Winning the Rally versus Scoring the Point

This article speculates that the benefits of competition may derive from the enjoyment and absorption of intrinsic competition, while the many negative effects of excessive competition, especially as seen in children and youth, may be due to extrinsic competition, the pursuit of wins for their own sake. Twelve practical suggestions are offered for physical education teachers and coaches who wish to bring intrinsic competition to the fore and de-emphasize extrinsic competition.

A temporal trophic shift from primary parasitism to facultative hyperparasitism during interspecific competition between two coevolved scelionid egg parasitoids.

Understanding competition between scelionid parasitoids that exploit the same host may provide insight into strategies that allow coexistence on a shared resource. Competition studies typically focus on interactions between native and exotic parasitoids that do not share an evolutionary history; however, coevolved parasitoids may be more likely to demonstrate strategies to avoid or exploit a shared resource. We examined intrinsic and extrinsic competition between Asian Trissolcus japonicus (Ashmead) and T. cultratus (Mayr) (Hymenoptera: Scelionidae) associated with Halyomorpha halys (Stål) (Hemiptera: Pentatomidae) that share an evolutionary history. Interspecific interactions were assessed by providing parasitized egg masses to each species at various intervals post‐parasitism, and measuring host acceptance, developmental suitability, and guarding behaviour. Trissolcus japonicus showed high acceptance of parasitized hosts up to 72 h following oviposition by T. cultratus, despite a very poor developmental outcome. In contrast, T. cultratus generally avoided ovipositing in H. halys eggs containing T. japonicus early‐instar larvae but did not avoid parasitizing H. halys that contained eggs and third instar larvae. The adaptive value of this behaviour was supported by developmental outcome: T. cultratus outcompeted T. japonicus eggs but not early‐instar larvae, and a trophic shift occurred wherein T. cultratus developed as a facultative hyperparasitoid on third instar T. japonicus larvae. Trissolcus japonicus guarded egg masses 8–12× longer and displayed more aggressive interactions than T. cultratus, suggesting T. japonicus is the superior extrinsic competitor. Development as a facultative hyperparasitoid provided a competitive niche for Asian T. cultratus and confirms its instrinsic competitive superiority. This also occurs in a biologically distinct European population of T. cultratus, suggesting that facultative hyperparasitism as a competitive strategy is retained in geographically separated populations that have not coevolved with H. halys or T. japonicus.

Extrinsic Inter- and Intraspecific Competition in Parasitoid Wasps

The diverse ecology of parasitoids is shaped by extrinsic competition, i.e., exploitative or interference competition among adult females and males for hosts and mates. Adult females use an array of morphological, chemical, and behavioral mechanisms to engage in competition that may be either intra- or interspecific. Weaker competitors are often excluded or, if they persist, use alternate host habitats, host developmental stages, or host species. Competition among adult males for mates is almost exclusively intraspecific and involves visual displays, chemical signals, and even physical combat. Extrinsic competition influences community structure through its role in competitive displacement and apparent competition. Finally, anthropogenic changes such as habitat loss and fragmentation, invasive species, pollutants, and climate change result in phenological mismatches and range expansions within host-parasitoid communities with consequent changes to the strength of competitive interactions. Such changes have important ramifications not only for the success of managed agroecosystems, but also for natural ecosystem functioning.

Interspecific Competition Between Idiobiont Larval Parasitoids of Emerald Ash Borer

New associations between parasitoid species is common in biological control with interactions ranging from coexistence to competitive exclusion. Spathius agrili and Spathius galinae, native to North East Asia, are two host-specific idiobiont larval parasitoids of the invasive emerald ash borer Agrilus planipennis that do not overlap in their native ranges but have been recently introduced to the United States for biocontrol of A. planipennis. We conducted laboratory experiments to determine the results of competition between these two species. Competition between parasitoid larvae on a single host (intrinsic competition) and competition between adult parasitoids for oviposition sites (extrinsic competition) were evaluated. Successful multiparasitism did not occur in any intrinsic trials. Extrinsic competition favored S. agrili, but S. galinae was not excluded. Competition lowered parasitism rates for both species, but overall parasitism was not lowered.

Host Patch Use and Potential Competitive Interactions Between Two Egg Parasitoids From the Family Scelionidae, Candidate Biological Control Agents of Bagrada hilaris (Hemiptera: Pentatomidae).

Two egg parasitoids from Pakistan, Gryon sp. nr. gonikopalense Sharma (Hymenoptera: Scelionidae) and Trissolcus hyalinipennis Rajmohana & Narendran (Hymenoptera: Scelionidae), are currently being investigated as potential classical biocontrol agents for Bagrada hilaris Burmeister. The former is the most promising because of its ability to attack B. hilaris eggs in soil, but the latter was recently discovered in California. This study detailed the patch use and oviposition behavior of both species and assessed their relative foraging efficiency. We also investigated possible competitive interactions by assessing 1) the occurrence of intra- and interspecific host discrimination, 2) mutual interference between females (extrinsic competition), 3) the outcome of multiparasitism (intrinsic competition), and 4) the effect of competition on host suppression. Our results showed that T. hyalinipennis females tended to probe repeatedly in the same hosts leading to longer host patch time and lower foraging efficiency. Both species avoided conspecific superparasitism, but not multiparasitism. When the two species foraged simultaneously, G. sp. nr. gonikopalense seemed to be slightly superior in extrinsic competition, while neither species seemed to have an advantage in intrinsic competition. Also, neither species could develop as a facultative hyperparasitoid, but they inflicted non-reproductive mortality on eggs containing 4-d-old larvae of their competitor. Lastly, host mortality inflicted by G. sp. nr. gonikopalense and T. hyalinipennis when acting together appeared to be additive. These results suggest that the presence of T. hyalinipennis in California is unlikely to reduce G. sp. nr. gonikopalense efficiency, but will instead improve the biological control of B. hilaris.

Interspecific competition between Peristenus spretus and Peristenus relictus (Hymenoptera: Braconidae), larval parasitoids of Apolygus lucorum (Hemiptera: Miridae)

Interspecific competition among different species of insect parasitoids may affect the establishment or efficacy of species used for biological control of a target pests. Peristenus spretus Chen & van Achterberg and Peristenus relictus Loan (syn. P. stygicus) (Hymenoptera: Braconidae) are two dominant solitary parasitoids of mirid bugs in China. In this study, interspecific competition between larvae of P. spretus and P. relictus for the nutrients inside their common host, the green plant bug (Apolygus lucorum Meyer-Dürin), as well as their extrinsic competition for host resources were investigated in the laboratory. We examined the effect of four different time intervals between oviposition by the two species (10 min, 24 h, 48 h, and 72 h) and two parasitoid sequences (i.e., P. spretus before P. relictus [Ps-Pr] and P. relictus before P. spretus (Pr-Ps)] in no-choice tests to measure nutrient competition between the two species. In choice tests, three densities of A. lucorum nymphs (40, 80, and 120) were presented to various combinations of P. spretus and P. relictus (1,1; 2,2; or 3,3 pairs). In addition, the developmental times of the immature stage (from egg to late instar larva) of P. spretus and P. relictus were determined. We found that neither the time interval between oviposition by the two competing species, nor the order the parasitoids attacked (Ps-Pr or Pr-Ps) had any influence on the percentage of parasitoid offspring that belonged a particular parasitoid species. Peristenus spretus dominated, accounting for 89.2–97.9% of parasitoid offspring in the no-choice nutrient competition experiment. Parasitoid sequence did, however, affect sex ratio in the dominant parasitoid’s offspring, with 48.2% of P. spretus offspring being female in the Pr-Ps sequence, while only 29.2% were female in the Ps-Pr sequence. Overall, in the host resource competition experiment, 64.7–82.2% of all emerged adult parasitoids of the offspring generation, for all host density treatments, were P. spretus. The percentage of emerged parasitoids that were a given species was significantly affected by host density, but neither by the number of pairs of P. spretus or P. relictus nor by the parasitoid density × host density interaction. The developmental time of the immature stage of P. spretus was 2.2 days shorter than that of P. relictus, an important factor in making P. spretus a superior competitor over P. relictus in all the competition trials. Based on these results, we conclude P. spretus will have an important advantage when competing with P. relictus, a factor that should be considered in any biological control program using these two parasitoids in China.

Interference competition between Aphidius matricariae and Praon volucre (Hymenoptera: Braconidae) attacking two common aphid species

ABSTRACTInterference competition is a common phenomenon that occurs among adult females of different species to gain the highest food resources at the same time. In this research, extrinsic competition between Aphidius matricariae and Praon volucre on different densities and stages of two important pests of greenhouse crops in the world, Aphis gossypii and Myzus persicae, were investigated. The results of this study showed that total percentage parasitism of second, third and fourth nymphal instars of A. gossypii and M. persicae by A. matricariae, and P. volucre were affected by extrinsic competition. A combination of A. matricariae and P. volucre on A. gossypii had a negative effect on performance of these parasitoids. Although extrinsic competition reduced the efficiency of A. matricariae and P. volucre on M. persicae, total parasitism of M. persicae by A. matricariae and P. volucre (combination of two parasitoids) increased compared to A. matricariae or P. volucre acting alone. Also the results indicated that the extrinsic competition between A. matricariae and P. volucre on both A. gossypii and M. persicae was apparently more intense when presented with 25 hosts compared to 50 and 100 hosts. The results of this research show important information to determine an appropriate combination of these two parasitoid wasps for biological control of A. gossypii and M. persicae in greenhouses.

Trade‐offs in parasitism efficiency and brood size mediate parasitoid coexistence, with implications for biological control of the invasive emerald ash borer

Summary Parasitoids often are selected for use as biological control agents because of their high host specificity, yet such host specificity can result in strong interspecific competition. Few studies have examined whether and how various extrinsic factors (such as parasitism efficiency, i.e. the ability to optimize host‐finding attack rates) influence the outcome of competition between parasitoids, even though they could have profound effects on the implementation of classical biological control programmes. To determine the potential influence of extrinsic competition and coexistence on host suppression efficacy, we compared parasitism by two larval parasitoids (Tetrastichus planipennisi and Spathius galinae) of the invasive emerald ash borer (EAB) Agrilus planipennis, under different host densities, parasitoid densities, host plant sizes and parasitoid–host ratios. Spathius galinae had significantly higher parasitism efficiency (≈4 times), but significantly lower brood size (>6 times) than that of T. planipennisi. The attack rates of hosts increased significantly with parasitoid density, whereas host density did not significantly affect multiparasitism. The parasitism rate of T. planipennisi on small host logs was significantly higher than that on large logs, while host plant (log) size had no significant impact on S. galinae parasitism. The multiparasitism rate was rather low regardless of host log size and parasitoid/host density, indicating that intrinsic competition between the two species of parasitoids might seldom occur in the field. The two species of parasitoids could therefore coexist in the same habitat, and any adverse effects on the suppression of EAB populations caused by competitive behaviour between the two species of parasitoids would likely be negligible. Synthesis and applications. Our findings suggest that introducing multiple species of parasitic natural enemies could be feasible for management of invasive species, but it is important to examine multiple extrinsic factors simultaneously when evaluating interspecific competition between them. Among these different extrinsic factors, we found that coexistence between parasitoids can be mediated by trade‐offs in their parasitism efficiency and brood sizes. Thus, the differences in life‐history traits of natural enemies could be used to select among biological control agents being considered for releases.

Interspecific extrinsic and intrinsic competitive interactions in egg parasitoids

Interspecific competitive interactions can occur either between adult parasitoids searching/exploiting hosts (extrinsic competition) or between parasitoid larvae developing within the same host (intrinsic competition). Understanding how interspecific competition between parasitoids can affect pest suppression is important for improving biological pest control. The purpose of this work was to review both extrinsic and intrinsic competition between egg parasitoid species. These are organisms that are often candidates for biological control programs due to their ability to kill the pest before the crop feeding stage. We first reviewed the literature about interspecific competitive abilities of adult parasitoids in terms of comparative host location strategies highlighting which ecological and behavioral factors are likely to shape extrinsic competition. Then we focused on the interspecific competitive interactions between immatures developing within the same host taking into account which factors play a key role in the outcome of intrinsic competition. Finally we conclude stressing on the need to elucidate the overall competitive interaction that parasitoid species may experience in the field in order to enhance biological control success.

Protein-imprinted materials: rational design, application and challenges

Protein imprinting is a promising tool for generating artificial biomimetic receptors with antibody-like specific recognition sites. Recently, protein-imprinted materials, as potential antibody substitutes, have attracted much attention in many fields, for example chemical sensors, chromatographic stationary phases, and artificial enzymes, owing to their long-term storage stability, potential re-usability, resistance to harsh environment, and low cost. In this critical review, we focus our discussion on the rational preparation of protein-imprinted materials in terms of choice of template, functional monomer, crosslinker, and polymerization format. In addition, several highlighted applications of protein-imprinted materials are emphasized, not only in well-known fields but also in some unique fields, for example proteomics and tissue engineering. Finally, we propose challenges arising from the intrinsic properties of protein imprinting, for example obtaining the template, heterogeneous binding, and extrinsic competition, for example immobilized aptamers.

Possible effects of inter-specific competition on the coexistence of two parasitoid species:Trichogramma brassicaeBezdenko andChelonus oculator(F.) (Hymenoptera: Trichogrammatidae, Braconidae)

Insect parasitoids have been widely studied, particularly due to their ecological implications through the study of the special relationships observed among this kind of species, as well as to their expression in mathematical models. However, there are still scarce studies on parasitoid relationships and their expression in more realistic mathematical models. The present work is aimed at deepening into competition relationships among parasitoids. Bearing this purpose in mind, the system shaped by two parasitoids was chosen: Trichogramma brassicae (idiobiont egg parasitoid) and Chelonus oculator (koinobiont egg-larval parasitoid). Both species compete against each other for the same host species (Lepidoptera). The results obtained in the laboratory point out that T. brassicae may be considered a better competitor than Ch. oculator. This is the result of the extrinsic competition due to the substances injected by the female during parasitization. However, our results show this classification into better and worse competitors inaccurate. Thus, these interspecific competition influences are detrimental to both parasitoid species. This is the first time that the effect of this competition is mentioned regarding parasitoid functional response. Our results and their ecological implications are reported and discussed.

Behavioral response of the egg parasitoid Ooencyrtus telenomicida to host-related chemical cues in a tritrophic perspective

The response of the generalist egg parasitoid Ooencyrtustelenomicida (Vassiliev) (Hymenoptera: Encyrtidae) to host-related chemical cues from tomato plants, Solanum lycopersicum L., and adults of Nezara viridula (L.) (Heteroptera: Pentatomidae) was investigated in laboratory-based no-choice and paired-choice tests. In Y-tube olfactometer experiments, when female wasps were exposed to volatiles from plants in different conditions, they were attracted only to volatiles produced by N. viridula adult-infested tomato plants. When female wasps were exposed to adults of N. viridula, they were attracted to volatiles from virgin males, and, at a lower level, to volatiles from mated females in preoviposition state. Finally, studies in open arena showed that chemical footprints left by adults of N. viridula did not induce arrestment responses in wasp females. These results are discussed in terms of extrinsic competition with other beneficial egg parasitoids that in field can compete for the same egg mass, since intraguild interactions may affect the success of a biological control program.

Inverse Hall–Petch relationship in the nanostructured TiO2: Skin-depth energy pinning versus surface preferential melting

The functional dependence of stress, elastic modulus, melting point, and their interdependence on the identities (bond order, nature, length, and strength) of a representative bond of the specimen has been established for deeper insight into the transition from the conventional Hall–Petch relationship (HPR) to the inverse HPR (IHPR) for nanostructured TiO2. Theoretical reproduction of the observed inverse HPR suggests that the intrinsic competition between the energy-density gain (elastic modulus enhancement) and the cohesive-energy remnant (melting point depression) in the grain boundaries originates and the extrinsic competition between the activation and the inhibition of atomic dislocations activates the IHPR.

Egg Size, Intrinsic Competition, and Lethal Interference in the Parasitoids Encarsia pergandiella and Encarsia formosa

Recent population dynamic theory predicts that disruption of biological control may occur when one parasitoid species' superiority in intrinsic competition is associated with a lower ability to find and exploit hosts (i.e., ability in extrinsic competition). One might expect such a trade-off, for instance, if parasitoids with larger (and fewer) eggs are more likely to prevail in intrinsic competition than species with smaller (and more numerous) eggs. We tested the idea that relative egg size could be used to predict the outcome of intrinsic competition in two closely related endoparasitoids, Encarsia pergandiella Howard and Encarsia formosa Gahan. Contrary to expectation, the parasitoid species with smaller eggs, E. pergandiella, prevailed in intrinsic competition, regardless of the order that hosts were exposed to the two species. In a literature survey, we found four studies of competing pairs of endoparasitoid species for which: (a) egg size estimates were available and (b) one species was consistently superior in intrinsic competition. In three of the four studies, the small-egged species prevailed in intrinsic competition, as we also found. Although E. formosa lost in intrinsic competition, this species negatively affected E. pergandiella's progeny production by host feeding on and killing hosts containing E. pergandiella eggs. E. formosa females also host fed on conspecific-parasitized hosts. As a mechanism of both intra- and interspecific interference competition, host feeding on parasitized hosts contradicts assumptions about the nature of interference competition in existing population dynamics models.

Intrinsic and extrinsic competitive interactions between two larval parasitoids of Heliothis virescens

Summary1. Competition between parasitoid species may be a key factor in the community dynamics of plant–herbivore‐parasitoid systems and is an important consideration in the selection and management of effective biological control agents.2. Interspecific competition can occur between adult parasitoids searching for hosts (extrinsic competition) and between multiple parasitoid larvae developing within a single host individual (intrinsic competition). A model system comprising the lepidopteran pest Heliothis virescens and two key hymenopteran endoparasitoids, Microplitis croceipes and Cardiochiles nigriceps, was employed to explore parasitoid host‐location strategies and the consequences of intrinsic and extrinsic competitive interactions between parasitoid species.3. The less specialised of the two parasitoids, M. croceipes, was found to have a shorter hatching time and to dominate intrinsic competition, except when its oviposition followed that of the more specialised parasitoid, C. nigriceps, by 16 h or more. This interval corresponded to the differential in hatching time between the two species.4. Cardiochiles nigriceps, however, displayed superior host‐searching efficiency that may compensate for its disadvantage in intrinsic competition. This parasitoid was more effective at detecting host infestation sites via airborne odours and at locating and attacking early instar host larvae than was M. croceipes.

An application of the state-trait concept to the need for achievement

This study concerned relationships between state (situationally defined) n Ach, trait n Ach, and sex of subject in 80 college freshmen. Three standard n Ach tests (the Adjective Check List, Personal Preference Schedule, and TAT measures) and the mean of n Ach states measured under nonarousal conditions were used as trait measures. State measures of n Ach were also obtained in four experimental conditions (Social Ability, Success-Failure, Conventional Arousal, and Extrinsic Competition). There were no significant correlations between the four trait measures. The mean of states in the neutral conditions was highly predictive of states in three of the four arousal conditions, but none of the standard trait tests predicted state arousal to any significant extent.