Functional biodiversity to improve pest control in organic cropping systems

A sustainable use of functional agrobiodiversity (FAB) providing habitats with suitable floral resources is needed to conserve and improve pest control by natural enemies in organic cropping and other low-input systems. We present an overview on our activities identifying appropriate flowering plants in relation to the antagonists-pest complex and quantifying benefits and limits in lab- and field tests. We have focussed on the control of apple-aphids and cabbage lepidopterean pests in two organic cropping systems. We can show that tailoring the flowering strips to the needs of specific natural enemies within a cropping system is a key issue for successful application of FAB. We found plants as Fagopyrum esculentum, Centaurea cyanus and Vicia sativa enhancing target parasitoids in cabbage, and we found Daucus carota, Carvum carvi, Pastinca sativa, Vicia sepium as promising plants in apple orchards. A successful on-farm application of FAB using flowering strips and companion plants is challenging and needs further analyses of its impact on pest control, considering different scales as plot, farm and landscape

Sperm competition affects sex allocation but not sperm morphology in a flatworm

Sperm competition has been shown to be an important evolutionary agent affecting the behaviour, physiology, and morphology of both males and females. One morphological trait that is particularly likely to be affected by sperm competition is sperm size because it is thought to influence the competitiveness of sperm by determining sperm longevity, motility, and/or their ability to displace competing sperm. Most comparative studies across taxa have found a positive relationship between the level of sperm competition and sperm length, but very few studies have tested for a phenotypically plastic adjustment of sperm morphology in response to sperm competition. In this study, we experimentally tested for an effect of sperm competition on phenotypic plasticity in sperm morphology in an obligately outcrossing simultaneous hermaphrodite, the free-living flatworm Macrostomum lignano, by either raising worms in monogamous pairs (no sperm competition) or in promiscuous groups (intense sperm competition). Worms in groups produced larger testes and smaller ovaries as predicted by sex allocation theory and as previously documented in this species. However, we found no evidence for an effect of group size on sperm morphology, measured as total sperm length, sperm body length, and the length of two different sperm appendages. We conclude that M. lignano may either be incapable of adjusting the sperm morphology in a phenotypically plastic way and/or that there might be no benefit of phenotypic plasticity in sperm traits in this specie

Strategic mating effort in a simultaneous hermaphrodite: The role of the partner's feeding status

Sexual selection theory for simultaneously hermaphroditic animals predicts an overall preference for inseminating partners that have a relatively higher female fecundity. Previous work on the link between male mating decisions and female fecundity has primarily focused on the effect of the partners' body size using existing variation in this trait within a study population. On the assumption that the body size is positively correlated with female fecundity, sperm donors should preferentially inseminate relatively larger individuals to obtain a higher fitness gain through their male sex function. However, empirical evidence for such size-dependent mate choice in simultaneous hermaphrodites is equivocal, possibly because of confounding variables. We studied the mating behavior of the simultaneously hermaphroditic flatworm Macrostomum lignano and tested for a strategic mating effort in response to the feeding status of the partner. We experimentally manipulated the feeding status of potential mating partners in order to generate variation in female fecundity among them and tested whether this affected the copulation number and the number of sperm that the focal worm managed to store in the partner's sperm storage organ. We found that the manipulation of the feeding status had a strong effect on the body size of the potential mating partners and that focal worms copulated more frequently with, and stored more sperm in well-fed partners compared to unfed partners. Our results suggest that M. lignano adjusts its mating effort in response to the feeding status of the mating partne

Determinants of female fecundity in a simultaneous hermaphrodite: the role of polyandry and food availability

Classical sexual selection theory assumes that the reproductive success of females is primarily limited by the resources available for egg production rather than by the number of mating partners. However, there is now accumulating evidence that multiple mating can entail fitness costs or benefits for females. In this study we investigated the effect of polyandry (i.e., the mating with different mating partners) and food availability on the reproductive output of the female sex function in an outcrossing simultaneous hermaphrodite, the free-living flatworm Macrostomum lignano. We exposed virgin worms to different group sizes, a treatment that has previously been shown to affect the level of polyandry in this species. Moreover, we manipulated the food availability throughout the subsequent egg laying period, during which the worms were kept in isolation. The number of offspring produced was used as an estimate of female fecundity. We found that food availability, but not group size, had a significant effect on female fecundity. Additionally, female fecundity was positively correlated with the number of stored sperm in the female sperm-storage organ at the time of isolation, but it was not correlated with body or ovary size of the worms. Our results suggest that female fecundity in M.lignano is primarily determined by the resources available for egg production, and not by the level of polyandry, confirming classic sexual selection theory for simultaneous hermaphrodite

Variation in sex allocation plasticity in three closely related flatworm species

Sex allocation (SA) theory for simultaneous hermaphrodites predicts an influence of group size on SA. Since group size can vary within an individual's lifetime, this can favor the evolution of phenotypically plastic SA. In an emerging comparative context, we here report on SA plasticity in three closely related Macrostomum flatworm species, namely Macrostomum janickei, Macrostomum cliftonensis, and Macrostomum mirumnovem. For each species, we experimentally raised worms in three group sizes (isolated, pairs, and octets) and two enclosure sizes (small and large) in all factorial combinations and studied the effects of these factors on different estimates of SA. In addition, we also evaluated whether isolated worms engage in self‐fertilization. We found that all species have plastic SA, with M. cliftonensis being more plastic than the other two species, as assessed by comparing standardized effect sizes of (a) the presence/absence of mating partners and (b) the strength of sexual competition. Moreover, we found that sperm production rate-but not sperm morphology-is plastic in M. cliftonensis, and that only M. mirumnovem self‐fertilized during our observation period. Our study suggests that both SA and SA plasticity can diverge even between closely related species

Evolution of testicular architecture in the Drosophilidae: A role for sperm length

Background: Evolutionary biologists have so far largely treated the testis as a black box with a certain size, a matching resource demand and a resulting sperm output. A better understanding of the way that the testis responds to selection may come from recent developments in theoretical biology aimed at understanding the factors that influence the evolution of tissue architecture (i.e. the logical organisation of a tissue). Here we perform a comparative analysis of aspects of testicular architecture of the fruit fly family Drosophilidae. Specifically, we collect published information on the number of first (or primary) spermatocytes in spermatogenesis, which allows to infer an important aspect of testicular architecture. Results: We show that testicular architecture is much more variable (both within and between species) than is generally appreciated. Moreover, the number of first spermatocytes is strongly correlated to the sperm length, which is inversely related to the sperm production, and thus the workload of the testis. Conclusion: Our study clearly documents that tissue architecture can evolve, and that in the Drosophilidae it may do so in response to sexual selection. We conclude that the testis of the Drosophilidae is a promising model organ to test recent models of tissue architecture

Sperm competition and the evolution of spermatogenesis

Spermatogenesis is a long and complex process that, despite the shared overall goal of producing the male gamete, displays striking amounts of interspecific diversity. In this review, we argue that sperm competition has been an important selection pressure acting on multiple aspects of spermatogenesis, causing variation in the number and morphology of sperm produced, and in the molecular and cellular processes by which this happens. We begin by reviewing the basic biology of spermatogenesis in some of the main animal model systems to illustrate this diversity, and then ask to what extent this variation arises from the evolutionary forces acting on spermatogenesis, most notably sperm competition. We explore five specific aspects of spermatogenesis from an evolutionary perspective, namely: (i) interspecific diversity in the number and morphology of sperm produced; (ii) the testicular organizations and stem cell systems used to produce them; (iii) the large number and high evolutionary rate of genes underpinning spermatogenesis; (iv) the repression of transcription during spermiogenesis and its link to the potential for haploid selection; and (v) the phenomenon of selection acting at the level of the germline. Overall we conclude that adopting an evolutionary perspective can shed light on many otherwise opaque features of spermatogenesis, and help to explain the diversity of ways in which males of different species perform this fundamentally important proces

Successful mating and hybridisation in two closely related flatworm species despite significant differences in reproductive morphology and behaviour

Reproductive traits are some of the fastest diverging characters and can serve as reproductive barriers. The free-living flatworm Macrostomum lignano, and its congener M. janickei are closely related, but differ substantially in their male intromittent organ (stylet) morphology. Here, we examine whether these morphological differences are accompanied by differences in behavioural traits, and whether these could represent barriers to successful mating and hybridization between the two species. Our data shows that the two species differ in many aspects of their mating behaviour. Despite these differences, the species mate readily with each other in heterospecific pairings. Although both species have similar fecundity in conspecific pairings, the heterospecific pairings revealed clear postmating barriers, as few heterospecific pairings produced F1 hybrids. These hybrids had a stylet morphology that was intermediate between that of the parental species, and they were fertile. Finally, using a mate choice experiment, we show that the nearly two-fold higher mating rate of M. lignano caused it to mate more with conspecifics, leading to assortative mating, while M. janickei ended up mating more with heterospecifics. Thus, while the two species can hybridize, the mating rate differences could possibly lead to higher fitness costs for M. janickei compared to M. lignano

Thraustochytrids as novel parasitic protists of marine free-living flatworms: Thraustochytrium caudivorum sp. nov. parasitizes Macrostomum lignano

The Labyrinthulomycota are a relatively poorly studied group of heterotrophic unicellular eukaryotes. They comprise three lineages, labyrinthulids, thraustochytrids, and aplanochytrids, which are all primarily marine organisms and considered to be important components of marine microbial communities. Recently a number of Labyrinthulomycota have been implicated as parasites of marine (but also terrestrial) plants and marine molluscs. Here we describe a new species of thraustochytrid, Thraustochytrium caudivorum sp. nov. that we have isolated from laboratory cultures of Macrostomum lignano (Rhabditophora, Macrostomorpha), a marine free-living flatworm. In these worms T. caudivorum can cause lesions, which start at the tip of the tail plate and which can lead to the dissolution of the posterior part of the animal. Although the worms can frequently cure these lesions and regenerate the lost parts, the lesions can also result in the complete dissolution of the animal. We describe this thraustochytrid based on pure agar cultures and infestations in the worm cultures. Moreover, we describe its pathological effects on the worms and its morphology using both light and electron microscopy. In addition, we report a phylogenetic analysis using a partial 18S rDNA sequence that allows us to place this new species within the thraustochytrids. Finally, we outline a protocol that allows to permanently remove the parasites from infested worm cultures. We conclude that thraustochytrids represent a novel group of parasites of free-living flatworm

Indirect genetic effects and sexual conflicts: partner genotype influences multiple morphological and behavioural reproductive traits in a flatworm

The expression of an individual's phenotypic traits can be influenced by genes expressed in its social partners. Theoretical models predict that such indirect genetic effects (IGEs) on reproductive traits should play an important role in determining the evolutionary outcome of sexual conflict. However, empirical tests of (i) whether reproductive IGEs exist, (ii) how they vary among genotypes, and (iii) whether they are uniform for different types of reproductive traits are largely lacking. We addressed this in a series of experiments in the simultaneously hermaphroditic flatworm Macrostomum lignano. We found strong evidence for IGEs on both morphological and behavioral reproductive traits. Partner genotype had a significant impact on the testis size of focal individuals—varying up to 2.4-fold—suggesting that IGEs could mediate sexual conflicts that target the male sex function. We also found that time to first copulation was affected by a genotype × genotype interaction between mating partners, and that partner genotype affected the propensity to copulate and perform the postcopulatory suck behavior, which may mediate conflicts over the fate of received ejaculate components. These findings provide clear empirical evidence for IGEs on multiple behavioral and morphological reproductive traits, which suggests that the evolutionary dynamics of these traits could be altered by genes contained in the social environment.PostprintPeer reviewe