When can host shifts produce congruent host and parasite phylogenies? A simulation approach

Congruence between host and parasite phylogenies is often taken as evidence for cospeciation. However, 'pseudocospeciation', resulting from host-switches followed by parasite speciation, may also generate congruent trees. To investigate this process and the conditions favouring its appearance, we here simulated the adaptive radiation of a parasite onto a new range of hosts. A very high congruence between the host tree and the resulting parasite trees was obtained when parasites switched between closely related hosts. Setting a shorter time lag for speciation after switches between distantly related hosts further increased the degree of congruence. The shape of the host tree, however, had a strong impact, as no congruence could be obtained when starting with highly unbalanced host trees. The strong congruences obtained were erroneously interpreted as the result of cospeciations by commonly used phylogenetic software packages despite the fact that all speciations resulted from host-switches in our model. These results highlight the importance of estimating the age of nodes in host and parasite phylogenies when testing for cospeciation and also demonstrate that the results obtained with software packages simulating evolutionary events must be interpreted with caution.     

Origin and domestication of the fungal wheat pathogen Mycosphaerella graminicola via sympatric speciation

The Fertile Crescent represents the center of origin and earliest known place of domestication for many cereal crops. During the transition from wild grasses to domesticated cereals, many host-specialized pathogen species are thought to have emerged. A sister population of the wheat-adapted pathogen Mycosphaerella graminicola was identified on wild grasses collected in northwest Iran. Isolates of this wild grass pathogen from 5 locations in Iran were compared with 123 M. graminicola isolates from the Middle East, Europe, and North America. DNA sequencing revealed a close phylogenetic relationship between the pathogen populations. To reconstruct the evolutionary history of M. graminicola, we sequenced 6 nuclear loci encompassing 464 polymorphic sites. Coalescence analyses indicated a relatively recent origin of M. graminicola, coinciding with the known domestication of wheat in the Fertile Crescent around 8,000-9,000 BC. The sympatric divergence of populations was accompanied by strong genetic differentiation. At the present time, no genetic exchange occurs between pathogen populations on wheat and wild grasses although we found evidence that gene flow may have occurred since genetic differentiation of the populations.     

Detecting the coevolution of biosequences--an example of RNA interaction prediction

A probabilistic graphical model is proposed in order to detect the coevolution between different sites in biological sequences. The model extends the continuous-time Markov process of sequence substitution for single nucleic or amino acids and imposes general constraints regarding simultaneous changes on the substitution rate matrix. Given a multiple sequence alignment for each molecule of interest and a phylogenetic tree, the model can predict potential interactions within or between nucleic acids and proteins. Initial validation of the model is carried out using tRNA and 16S rRNA sequence data. The model accurately identifies the secondary interactions of tRNA as well as several known tertiary interactions. In addition, results on 16S rRNA data indicate this general and simple coevolutionary model outperforms several other parametric and nonparametric methods in predicting secondary interactions. Furthermore, the majority of the putative predictions exhibit either direct contact or proximity of the nucleotide pairs in the 3-dimensional structure of the Thermus thermophilus ribosomal small subunit. The results on RNA data suggest a general model of coevolution might be applied to other types of interactions between protein, DNA, and RNA molecules.     

Functional coevolutionary networks of the Hsp70-Hop-Hsp90 system revealed through computational analyses

Currently, the identification of groups of amino acid residues that are important in the function, structure, or interaction of a protein can be both costly and prohibitively complex, involving vast numbers of mutagenesis experiments. Here, we present the application of a novel computational method, which identifies the presence of coevolution in a data set, thereby enabling the a priori identification of amino acid residues that play an important role in protein function. We have applied this method to the heat shock protein (Hsp) protein-folding system, studying the network between Hsp70, Hsp90, and Hop (heat shock-organizing protein). Our analysis has identified functional residues within the tetratricopeptide repeat (TPR) 1 and 2A domains in Hop, previously shown to be interacting with Hsp70 and Hsp90, respectively. Further, we have identified significant residues elsewhere in Hop within domains that have been recently proposed as being important for Hop interaction with Hsp70 and/or Hsp90. In addition, several amino acid sites present in groups of coevolution were identified as 3-dimensionally or linearly proximal to functionally important sites or domains. Based on our results, we also investigate a further functional domain within Hop, between TPR1 and TPR2A, which we suggest as being functionally important in the interaction of Hop with both Hsp70 and Hsp90 whether directly or otherwise. Our method has identified all the previously characterized functionally important regions in this system, thereby indicating the power of this method in the a priori identification of important regions for site-directed mutagenesis studies.     

The correlation between immunocompetence and an ornament trait changes over lifetime in Panorpa vulgaris scorpionflies

The immunocompetence-handicap hypothesis posits that costly male ornament traits might function to signal superior heritable immunocompetence to females. Quite a number of studies have aimed at testing this hypothesis. Yet the empirical data obtained so far are ambiguous. Many studies analysed the phenotypic correlation between handicap expression and immunocompetence at the same time point. However, since immunocompetence may change drastically over an individual's lifetime, such a singular measurement may not represent genetic differences among males and the benefits of choosing handicapped males for females might thus be weak. Here, I tested the correlation of a potential immunocompetence-handicap, the production of salivary secretions as nuptial gifts in a scorpionfly (Panorpa vulgaris), with immunocompetence at two different time points. I found a positive correlation with the handicap, but only if immunocompetence was measured shortly after expression of the handicap, i.e. briefly after mating in 2 weeks old scorpionflies. By contrast, there was no correlation with immunocompetence of the same flies at a younger age, i.e. shortly after adult emergence and a weak, insignificant trend for increased immunocompetence in offspring. These results are in agreement with positive phenotypic correlations between immunocompetence and handicap expression reported from other species, but advise caution when generalizing such one-time correlations.     

Rapid evolution and ecological host-parasite dynamics

Traditionally, the termination of parasite epidemics has been attributed to ecological causes: namely, the depletion of susceptible hosts as a result of mortality or acquired immunity. Here, we suggest that epidemics can also end because of rapid host evolution. Focusing on a particular host–parasite system, Daphnia dentifera and its parasite Metschnikowia bicuspidata , we show that Daphnia from lakes with recent epidemics were more resistant to infection and had less variance in susceptibility than Daphnia from lakes without recent epidemics. However, our studies revealed little evidence for genetic variation in infectivity or virulence in Metschnikowia . Incorporating the observed genetic variation in host susceptibility into an epidemiological model parameterized for this system reveals that rapid evolution can explain the termination of epidemics on time scales matching what occurs in lake populations. Thus, not only does our study provide rare evidence for parasite-mediated selection in natural populations, it also suggests that rapid evolution has important effects on short-term host–parasite dynamics.     

A meta-analysis of the interactive effects of UV and drought on plants

Interactions between climate change and UV penetration in the biosphere are resulting in the exposure of plants to new combinations of UV radiation and drought. In theory, the impacts of combinations of UV and drought may be additive, synergistic or antagonistic. Lack of understanding of the impacts of combined treatments creates substantial uncertainties that hamper predictions of future ecological change. Here, we compiled information from 52 publications and analysed the relative impacts of UV and/or drought. Both UV and drought have substantial negative effects on biomass accumulation, plant height, photosynthesis, leaf area and stomatal conductance and transpiration, while increasing stress-associated symptoms such as MDA accumulation and reactive-oxygen-species content. Contents of proline, flavonoids, antioxidants and anthocyanins, associated with plant acclimation, are upregulated both under enhanced UV and drought. In plants exposed to both UV and drought, increases in plant defense responses are less-than-additive, and so are the damage and growth retardation. Less-than-additive effects were observed across field, glasshouse and growth-chamber studies, indicating similar physiological response mechanisms. Induction of a degree of cross-resistance seems the most likely interpretation of the observed less-than-additive responses. The data show that in future climates, the impacts of increases in drought exposure may be lessened by naturally high UV regimes.     

An out-of-body experience: the extracellular dimension for the transmission of mutualistic bacteria in insects

Across animals and plants, numerous metabolic and defensive adaptations are a direct consequence of symbiotic associations with beneficial microbes. Explaining how these partnerships are maintained through evolutionary time remains one of the central challenges within the field of symbiosis research. While genome erosion and co-cladogenesis with the host are well-established features of symbionts exhibiting intracellular localization and transmission, the ecological and evolutionary consequences of an extracellular lifestyle have received little attention, despite a demonstrated prevalence and functional importance across many host taxa. Using insect–bacteria symbioses as a model, we highlight the diverse routes of extracellular symbiont transfer. Extracellular transmission routes are unified by the common ability of the bacterial partners to survive outside their hosts, thereby imposing different genomic, metabolic and morphological constraints than would be expected from a strictly intracellular lifestyle. We emphasize that the evolutionary implications of symbiont transmission routes (intracellular versus extracellular) do not necessarily correspond to those of the transmission mode (vertical versus horizontal), a distinction of vital significance when addressing the genomic and physiological consequences for both host and symbiont.     

Geographical variation in mutualistic networks: similarity,turnover and partner fidelity

Although species and their interactions in unison represent biodiversity and all the ecological and evolutionary processes associated with life, biotic interactions have, contrary to species, rarely been integrated into the concepts of spatial β-diversity. Here, we examine β-diversity of ecological networks by using pollination networks sampled across the Canary Islands. We show that adjacent and distant communities are more and less similar, respectively, in their composition of plants, pollinators and interactions than expected from random distributions. We further show that replacement of species is the major driver of interaction turnover and that this contribution increases with distance. Finally, we quantify that species-specific partner compositions (here called partner fidelity) deviate from random partner use, but vary as a result of ecological and geographical variables. In particular, breakdown of partner fidelity was facilitated by increasing geographical distance, changing abundances and changing linkage levels, but was not related to the geographical distribution of the species. This highlights the importance of space when comparing communities of interacting species and may stimulate a rethinking of the spatial interpretation of interaction networks. Moreover, geographical interaction dynamics and its causes are important in our efforts to anticipate effects of large-scale changes, such as anthropogenic disturbances.