On fitness in structured metapopulations

In this paper we derive a general expression measuring fitness in general structured metapopulation models. We apply the theory to a model structured by local population size and in which local dynamics is explicitly modelled. In particular, we calculate the evolutionarily stable dispersal strategy for individuals that can assess the local population density in the case where only dispersal is subject to evolutionary control but all other model ingredients are assumed fixed. We show that there exists a threshold size such that at ESS everyone should stay as long as the population size is below the threshold and everyone should disperse immediately as the population size reaches the threshold. Received: 13 August 1999 / Revised version: 2 May 2001 / Published online: 12 October 2001     

Costs,benefits, and loss of vertically transmitted symbionts affect host population dynamics

The costs and benefits of symbiotic interactions may vary with host and symbiont ontogeny. Effects of symbionts at different stages of host development or on different host demographic rates do not contribute equally to fitness. Although rarely applied, a population dynamics approach that integrates over the host life cycle is therefore necessary for capturing the net costs or benefits and, thus, the mutualistic or parasitic nature of symbioses. Using the native, disturbance‐specialist grass Agrostis hyemalis, we asked how a symbiotic endophyte affected the population dynamics of its host and how imperfect vertical transmission influenced symbiont frequency in a late successional environment. A size‐structured integral projection model (IPM) parameterized with experimental field data showed that greater rates of individual growth and reproduction for endophyte‐symbiotic (E+) hosts outweighed their lower rates of survival, leading to a net positive effect of symbiosis on equilibrium plant population growth (slower rate of extinction). Given that populations under going successional transitions are unlikely to be at an equilibrium size structure, we also conducted transient analysis that showed an initial short‐term cost to endophyte symbiosis. We used a megamatrix approach to link E? and E+ IPMs via imperfect vertical transmission and found that this parameter strongly influenced the frequency of symbiosis via complex interactions with host demographic rates. Overall, our population dynamics approach improves the ability to characterize the outcome of symbiotic interactions, and results suggest that particular attention should be paid to interactions between the rate of vertical transmission and host demography.     

Evolution of specialization in resource utilization in structured metapopulations

We study the evolution of resource utilization in a structured discrete-time metapopulation model with an infinite number of patches, prone to local catastrophes. The consumer faces a trade-off in the abilities to consume two resources available in different amounts in each patch. We analyse how the evolution of specialization in the utilization of the resources is affected by different ecological factors: migration, local growth, local catastrophes, forms of the trade-off and distribution of the resources in the patches. Our modelling approach offers a natural way to include more than two patch types into the models. This has not been usually possible in the previous spatially heterogeneous models focusing on the evolution of specialization.     

Evolutionary suicide and evolution of dispersal in structured metapopulations

 We study the evolution of dispersal in a structured metapopulation model. The metapopulation consists of a large (infinite) number of local populations living in patches of habitable environment. Dispersal between patches is modelled by a disperser pool and individuals in transit between patches are exposed to a risk of mortality. Occasionally, local catastrophes eradicate a local population: all individuals in the affected patch die, yet the patch remains habitable. We prove that, in the absence of catastrophes, the strategy not to migrate is evolutionarily stable. Under a given set of environmental conditions, a metapopulation may be viable and yet selection may favor dispersal rates that drive the metapopulation to extinction. This phenomenon is known as evolutionary suicide. We show that in our model evolutionary suicide can occur for catastrophe rates that increase with decreasing local population size. Evolutionary suicide can also happen for constant catastrophe rates, if local growth within patches shows an Allee effect. We study the evolutionary bifurcation towards evolutionary suicide and show that a discontinuous transition to extinction is a necessary condition for evolutionary suicide to occur. In other words, if population size smoothly approaches zero at a boundary of viability in parameter space, this boundary is evolutionarily repelling and no suicide can occur. Received: 10 November 2000 / Revised version: 13 February 2002 / Published online: 17 July 2002     

Joint evolution of specialization and dispersal in structured metapopulations

We study the joint evolution of dispersal and specialization concerning resource usage in a mechanistically underpinned structured discrete-time metapopulation model. We show that dispersal significantly affects the evolution of specialization and that specialization is a key factor that determines the possibility of evolutionary branching in dispersal propensity. Allowing both dispersal propensity and specialization to evolve as a consequence of natural selection is necessary in order to understand the evolutionary dynamics. The joint evolution of dispersal and specialization forms a natural evolutionary path leading to the coexistence of generalists and specialists. We show that in this process, the number of different patch types and the resource distribution are essential.     

Evolution of specialization in resource utilization in structured metapopulations

We study the evolution of resource utilization in a structured discrete-time metapopulation model with an infinite number of patches, prone to local catastrophes. The consumer faces a trade-off in the abilities to consume two resources available in different amounts in each patch. We analyse how the evolution of specialization in the utilization of the resources is affected by different ecological factors: migration, local growth, local catastrophes, forms of the trade-off and distribution of the resources in the patches. Our modelling approach offers a natural way to include more than two patch types into the models. This has not been usually possible in the previous spatially heterogeneous models focusing on the evolution of specialization.     

On the evolution of specialization with a mechanistic underpinning in structured metapopulations

We analyze the evolution of specialization in resource utilization in a discrete-time metapopulation model using the adaptive dynamics approach. The local dynamics in the metapopulation are based on the Beverton-Holt model with mechanistic underpinnings. The consumer faces a trade-off in the abilities to consume two resources that are spatially heterogeneously distributed to patches that are prone to local catastrophes. We explore the factors favoring the spread of generalist or specialist strategies. Increasing fecundity or decreasing catastrophe probability favors the spread of the generalist strategy and increasing environmental heterogeneity enlarges the parameter domain where the evolutionary branching is possible. When there are no catastrophes, increasing emigration diminishes the parameter domain where the evolutionary branching may occur. Otherwise, the effect of emigration on evolutionary dynamics is non-monotonous: both small and large values of emigration probability favor the spread of the specialist strategies whereas the parameter domain where evolutionary branching may occur is largest when the emigration probability has intermediate values. We compare how different forms of spatial heterogeneity and different models of local growth affect the evolutionary dynamics. We show that even small changes in the resource dynamics may have outstanding evolutionary effects to the consumers.     

Many corals host thermally resistant symbionts in high-temperature habitat

Physiologically distinct lines of dinoflagellate symbionts, Symbiodinium spp., may confer distinct thermal tolerance thresholds on their host corals. Therefore, if a coral can alternately host distinct symbionts, changes in their Symbiodinium communities might allow corals to better tolerate increasing environmental temperatures. However, researchers are currently debating how commonly coral species can host different symbiont types. We sequenced chloroplast 23 s rDNA from the Symbiodinium communities of nine reef-building coral species across two thermally distinct lagoon pools separated by ~500 m. The hotter of these pools reaches 35°C in the summer months, while the other pool’s maximum temperature is 1.5°C cooler. Across 217 samples from nine species, we found a single haplotype in both Symbiodinium clades A and D, but four haplotypes in Symbiodinium clade C. Eight of nine species hosted a putatively thermally resistant member of clade D Symbiodinium at least once, one of which hosted this clade D symbiont exclusively. Of the remaining seven that hosted multiple Symbiodinium types, six species showed higher proportions of the clade D symbiont in the hotter pool. Average percentage rise in the frequency of the clade D symbiont from the hotter to cooler pool was 52% across these six species. Even though corals hosted members of both the genetically divergent clades D and C Symbiodinium, some showed patterns of host–symbiont specificity within clade C. Both Acropora species that hosted clade C exclusively hosted a member of sub-clade C2, while all three Pocillopora species hosted a member of sub-clade C1 (sensu van Oppen et al. 2001). Our results suggest that coral–algal symbioses often conform to particular temperature environments through changes in the identity of the algal symbiont.     

Transformed bacterial symbionts re-introduced to and detected in host gut

Strains of Enterobacter agglomerans and Klebsiella pneumoniae isolated from Rhagoletis completa Cresson were engineered to express transgenic fluorescent proteins (ECFP, DsRed). These bacteria were introduced into flies by feeding the flies a sucrose solution in which the bacteria were suspended. The transgenic and heterologous marker protein was expressed and visible in the bacteria after they were ingested by WHF and while they were in the fly gut. We describe the plasmids used to transform these bacteria and demonstrate expression of heterologous proteins from the transforming plasmids and discuss the implications for future pest control strategies. Received: 14 September 2001 / Accepted: 22 October 2001     

Standing genetic variation in host preference for mutualist microbial symbionts

Many models of mutualisms show that mutualisms are unstable if hosts lack mechanisms enabling preferential associations with mutualistic symbiotic partners over exploitative partners. Despite the theoretical importance of mutualism-stabilizing mechanisms, we have little empirical evidence to infer their evolutionary dynamics in response to exploitation by non-beneficial partners. Using a model mutualism—the interaction between legumes and nitrogen-fixing soil symbionts—we tested for quantitative genetic variation in plant responses to mutualistic and exploitative symbiotic rhizobia in controlled greenhouse conditions. We found significant broad-sense heritability in a legume host''s preferential association with mutualistic over exploitative symbionts and selection to reduce frequency of associations with exploitative partners. We failed to detect evidence that selection will favour the loss of mutualism-stabilizing mechanisms in the absence of exploitation, as we found no evidence for a fitness cost to the host trait or indirect selection on genetically correlated traits. Our results show that genetic variation in the ability to preferentially reduce associations with an exploitative partner exists within mutualisms and is under selection, indicating that micro-evolutionary responses in mutualism-stabilizing traits in the face of rapidly evolving mutualistic and exploitative symbiotic bacteria can occur in natural host populations.     

Periodic local disturbance in host-parasitoid metapopulations: host suppression and parasitoid persistence

Within many agricultural systems, insect pests and their natural enemies are forced to persist as a metapopulation, continuously recolonizing patches following disturbance through harvesting or spraying with insecticides. Despite the need to understand factors influencing biocontrol success, few theoretical studies of host-parasitoid interactions have examined the potential impact of local disturbance within a metapopulation framework. Here, we add periodic local mortality to series of classical host-parasitoid models to examine its effect on host suppression and parasitoid persistence. Using a deterministic lattice model, we show that despite the wide range of complex dynamics generated at the patch level, the region wide pattern of disturbance is the key factor influencing host suppression. The level of host suppression achieved can be understood in terms of both the strength of density dependent parasitism, and the relative amounts of host and parasitoid mixing amongst patches of different ages. Local dispersal among patches is sufficient to ensure coexistence of the host and parasitoid, though persistence is not necessarily associated with the formation of self-organized spatial structures reported in previous studies. Finally, a stochastic version of the model is developed, in order to highlight how the effects of demographic stochasticity may influence biocontrol success in highly disturbed agricultural systems.     

Facultative symbionts are associated with host plant specialization in pea aphid populations

The pea aphid, Acyrthosiphon pisum, shows significant reproductive isolation and host plant specialization between populations on alfalfa and clover in New York. We examine whether specialization is seen in pea aphids in California, and whether fitness on alternative host plants is associated with the presence of bacterial symbionts. We measured the fitness of alfalfa- and clover-derived aphids on both types of plants and found no evidence for specialization when all aphid lineages were considered simultaneously. We then screened all aphids for the presence of four facultative bacterial symbionts: PAR, PASS, PABS and PAUS. Aphids with PAUS were host-plant specialized, having twice as many offspring as other aphids on clover, and dying on alfalfa. Other aphids showed no evidence of specialization. Additionally, aphids with PABS had 50% more offspring than aphids with PASS when on alfalfa. Thus, specialist and generalist aphid lineages coexist, and specialization is symbiont associated. Further work will resolve whether PAUS is directly responsible for this variation in fitness or whether PAUS is incidentally associated with host-plant specialized aphid lineages.     

Effects of bacterial secondary symbionts on host plant use in pea aphids

Aphids possess several facultative bacterial symbionts that have important effects on their hosts'' biology. These have been most closely studied in the pea aphid (Acyrthosiphon pisum), a species that feeds on multiple host plants. Whether secondary symbionts influence host plant utilization is unclear. We report the fitness consequences of introducing different strains of the symbiont Hamiltonella defensa into three aphid clones collected on Lathyrus pratensis that naturally lack symbionts, and of removing symbionts from 20 natural aphid–bacterial associations. Infection decreased fitness on Lathyrus but not on Vicia faba, a plant on which most pea aphids readily feed. This may explain the unusually low prevalence of symbionts in aphids collected on Lathyrus. There was no effect of presence of symbiont on performance of the aphids on the host plants of the clones from which the H. defensa strains were isolated. Removing the symbiont from natural aphid–bacterial associations led to an average approximate 20 per cent reduction in fecundity, both on the natural host plant and on V. faba, suggesting general rather than plant-species-specific effects of the symbiont. Throughout, we find significant genetic variation among aphid clones. The results provide no evidence that secondary symbionts have a major direct role in facilitating aphid utilization of particular host plant species.     

Vertically transmitted fungal endophytes: different responses of host-parasite systems to environmental conditions

The relationship between vertically transmitted asexual fungal grass endophytes and their hosts is considered to be mutualistic. Results from agronomic field support this line of reasoning but recent studies have shown more variable results in natural systems. We investigated how high and low nutrient and water treatments affected biomass allocation patterns of endophyte‐infected and uninfected Festuca pratensis and F. rubra in greenhouse experiments over two growing seasons. Irrespective of infection status, both grass species showed improved performance on highly fertilized and watered soils. However, infected F. pratensis plants produced larger tillers than endophyte‐free plants on soil low in nutrients and water in the first growing season, although they (E+) otherwise showed decreased performance on nutrient‐poor soil. In low nutrient and water conditions, endophyte‐infected plants produced less tillers and had lower total biomass compared to uninfected plants, and displayed a negative phenotypic correlation between seed production and vegetative growth. The latter indicates costs of reproduction when the plant shares common resources with the fungal endophyte. However, endophyte infection status (E+, E?) interacted significantly with the soil fertilisation in terms of plant growth, having a stronger positive effect on growth in infected F. pratensis plants. In F. rubra, endophyte‐infected plants showed higher vegetative growth in fertilized and watered soils compared to uninfected plants. However, infected plants tended to produce fewer inflorescences. This had no effect on seed production, perhaps because seed production was partly replaced by asexual pseudovivipary. Contrary to the general assumption in the literature that fungal endophytes are plant mutualists, these findings suggest that the costs of endophytes may outweigh their benefits in resource limited conditions. However, the costs of endophyte infections appear to differ among the grass species studied; costs of endophytes were mainly detected in F. pratensis under low nutrient conditions. We propose that differences in response to endophyte infection in these species may depend on the differences in life‐history strategies and environmental requirements of these two fescue and fungal species and may change during the life span of the plant.     

Zooxanthellar symbionts shape host sponge trophic status through translocation of carbon

Sponges belonging to the genus Cliona are common inhabitants of many coral reefs, and as bioeroders, they play an important role in the carbonate cycle of the reef. Several Cliona species maintain intracellular populations of dinoflagellate zooxanthellae (i.e., Symbiodinium spp.), which also form symbioses with a variety of other invertebrates and protists (e.g., corals, molluscs, foraminifera). Unlike the case of coral symbioses, however, almost nothing is known of the metabolic interaction between sponges and their zooxanthella symbionts. To assess this interaction, we performed a tracer experiment to follow C and N in the system, performed a reciprocal transplant experiment, and measured the stable carbon isotope ratio of Cliona spp. with and without zooxanthellae to study the influence of environment on the interaction. We found strong evidence of a transfer of C from zooxanthellae to their sponge hosts but no evidence of a transfer of N from sponge to zooxanthellae. We also saw significant influences of the environment on the metabolism of the sponges. Finally, we observed significant differences in carbon metabolism of sponge species with and without symbionts. These data strongly support hypotheses of metabolic integration between zooxanthellae and their sponge host and extend our understanding of basic aspects of benthic-pelagic coupling in shallow-water marine environments.     

Pathogen invasion and host extinction in lattice structured populations

We examined the propagation of an infectious disease and the eventual extinction of the host population in a lattice-structured population. Both the host colonization and pathogen transmission processes are assumed to be restricted to act between the nearest neighbor sites. The model is analyzed by an improved version of pair approximation (IPA). Pair approximation is a technique to trace the dynamics of the number of nearest neighbor pairs having particular states, and IPA takes account of the clustering property of lattice models more precisely. The results are checked by computer simulations. The analysis shows: (i) in a one-dimensional lattice population, a pathogen cannot invade a host population no matter how large is the transmission rate; (ii) in a two-dimensional lattice population, pathogens will drive the host to extinction if the transmission rate is larger than a threshold. These results indicate that spatially structured population models may give qualitatively different results from conventional population models, such as Lotka-Volterra ones, without spatial structure.     

Vertically structured prokaryotic community can control the efficiency of the biological pump in the oceans

Despite recent advances in molecular and metagenomic approaches, it is still unclear how spatiotemporal variations in microbial communities influence the biological pump, exporting organic carbon from the surface to the deep oceans. We address this important problem by constructing a simple model of a prokaryotic metacommunity in which two generalist ecotypes compete for two resources. The first and the second ecotypes have, respectively, a higher preference for particulate organic carbon (POC) and dissolved organic carbon. Sinking of POC would lead to a higher abundance of the first ecotype in the deep layer compared to the surface layer, but vertical mixing in turn generates a net upward flux of this ecotype to the surface layer. This upward movement accelerates the shifts in the community composition during the phytoplankton bloom, contributing to a higher efficiency in POC remineralization at the surface layer and reducing the carbon flux to the deep layer.