The size and shape of parasitic larvae of naiads (Unionidae) are not dependent on female size

The naiads, large freshwater mussels (Unionida), have very long life spans, are large-bodied, and produce thousands to millions of larvae (glochidia) which typically must attach to host fish tissues to metamorphose into a juvenile mussel. Glochidia develop within a female's marsupial gill demibranch, thus their number is restricted by female size. However, larger mussels acquire more energy, which could be invested in either larger-sized glochidia, in a more glochidia, or a combination of both. The high level of host specialization seen in many naiads may constrain glochidial size and shape around a narrow optimum, while naiads that use a wide range of host fishes may be predicted to possess greater plasticity in glochidial morphology. In this paper, we investigated the relationship between maternal body size and progeny body size and shape, aided by modern digital microscopy. We analyzed the between- and within- species variation of glochidia size and shape relative to female size in four widespread species of European naiads: Anodonta anatina, Anodonta cygnea, Unio crassus and Unio tumidus. Whereas the total reproductive output is collinear with female body size, substantial differences between species in glochidia size were found within genus Anodonta, but not genus Unio where glochidial size is remarkably consistent. The glochidial shape, however, differed within both Unio and Anodonta. We interpret this constant within-species glochidial size in Unio as reflecting a constraint imposed by the likelihood of successful transmission onto and off from a narrow range of hosts, whereas their shape seems to be less constrained. The Anodonta species, inhabiting a wide spectrum of habitats and using more than twice the number of fish hosts than Unio spp., have larger glochidia with greater variation in size and shape. Our results suggest that measures of glochidial variability may also serve as an indicator of host specificity in other naiads.

The classification of dwarf mistletoes (Arceuthobium spp., Viscaceae) in section Campylopoda, series Campylopoda

The taxonomic classification of dwarf mistletoes (Arceuthobium spp., Viscaceae) is complicated due to their reduced morphology, requiring the integration of not only morphology but also phenology, geography, and host relationships. This has been particularly true for the classification of taxa in subgenus Vaginata, section Campylopoda, series Campylopoda. Most of the species in this group have been recently circumscribed in synonymy with or reduced to subspecies of Arceuthobium campylopodum Engelm; however, we contend they deserve separate species recognition. To address this question, we have conducted morphological analyses of the taxa in ser. Campylopoda using univariate and multivariate statistical analyses. Our results have demonstrated that these taxa can be determined to species using morphological data without consideration of geographic location or host specificity; however, the host specialization and geographic distribution exhibited by these taxa also supports their classification as species. Here, we discuss the evidence supporting the specific classification of ser. Campylopoda taxa. This taxonomic framework permits the treatment of several dwarf mistletoe populations with geographically restricted distributions, fewer morphological differences, and specialized host affinities as subspecies of Arceuthobium abietinum (Engelm.) Abrams, Arceuthobium microcarpum (Engelm.) Hawksw. & Wiens, and Arceuthobium tsugense (Rosend.) G.N.Jones. It is also the most practical classification for the management of these economically and ecologically important parasitic plants.

A single amino acid polymorphism in a conserved effector of the multihost blast fungus pathogen expands host-target binding spectrum

Accelerated gene evolution is a hallmark of pathogen adaptation and specialization following host-jumps. However, the molecular processes associated with adaptive evolution between host-specific lineages of a multihost plant pathogen remain poorly understood. In the blast fungus Magnaporthe oryzae (Syn. Pyricularia oryzae), host specialization on different grass hosts is generally associated with dynamic patterns of gain and loss of virulence effector genes that tend to define the distinct genetic lineages of this pathogen. Here, we unravelled the biochemical and structural basis of adaptive evolution of APikL2, an exceptionally conserved paralog of the well-studied rice-lineage specific effector AVR-Pik. Whereas AVR-Pik and other members of the six-gene AVR-Pik family show specific patterns of presence/absence polymorphisms between grass-specific lineages of M. oryzae, APikL2 stands out by being ubiquitously present in all blast fungus lineages from 13 different host species. Using biochemical, biophysical and structural biology methods, we show that a single aspartate to asparagine polymorphism expands the binding spectrum of APikL2 to host proteins of the heavy-metal associated (HMA) domain family. This mutation maps to one of the APikL2-HMA binding interfaces and contributes to an altered hydrogen-bonding network. By combining phylogenetic ancestral reconstruction with an analysis of the structural consequences of allelic diversification, we revealed a common mechanism of effector specialization in the AVR-Pik/APikL2 family that involves two major HMA-binding interfaces. Together, our findings provide a detailed molecular evolution and structural biology framework for diversification and adaptation of a fungal pathogen effector family following host-jumps.

The Evolution of Host Specialization in an Insect Pathogen

Niche breadth coevolution between biotic partners underpins theories of diversity and co-existence and influences patterns of disease emergence and transmission in host-parasite systems. Despite these broad implications, we still do not fully understand how the breadth of parasites’ infectivity evolves, the nature of any associated costs, or the genetic basis of specialization. Here, we serially passage a granulosis virus on multiple inbred populations of its Plodia interpunctella host to explore the dynamics and outcomes of specialization. In particular, we collect time series of phenotypic and genetic data to explore the dynamics of host genotype specialization throughout the course of experimental evolution and examine two fitness components. We find that the Plodia interpunctella granulosis virus consistently evolves increases in overall specialization, but that our two fitness components evolve independently such that lines specialize in either productivity or infectivity. Furthermore, we find that specialization in our experiment is a highly polygenic trait best explained by a combination of evolutionary mechanisms including conditionally positive fitness asymmetries and mutation accumulation. These results are important for understanding the evolution of specialization in host-parasite interactions and its broader implications for co-existence, diversification, and infectious disease management.

Defensive Symbionts and the Evolution of Parasitoid Host Specialization

Insect host–parasitoid interactions abound in nature and are characterized by a high degree of host specialization. In addition to their behavioral and immune defenses, many host species rely on heritable bacterial endosymbionts for defense against parasitoids. Studies on aphids and flies show that resistance conferred by symbionts can be very strong and highly specific, possibly as a result of variation in symbiont-produced toxins. I argue that defensive symbionts are therefore an important source of diversifying selection, promoting the evolution of host specialization by parasitoids. This is likely to affect the structure of host–parasitoid food webs. I consider potential changes in terms of food web complexity, although the nature of these effects will also be influenced by whether maternally transmitted symbionts have some capacity for lateral transfer. This is discussed in the light of available evidence for horizontal transmission routes. Finally, I propose that defensive mutualisms other than microbial endosymbionts may also exert diversifying selection on insect parasitoids. Expected final online publication date for the Annual Review of Entomology, Volume 67 is January 2022. Please see http://www.annualreviews.org/page/journal/pubdates for revised estimates.

Taxonomic and pathogenic diversity of the blast pathogen populations infecting wheat and grasses in Minas Gerais

The blast disease of Poaceae is caused by a large species complex, among which P. oryzae is composed of several host-specialized lineages. The Pyricularia oryzae Triticum pathotype (PoT) causes the blast disease in wheat, but is also capable of infecting other grasses, which may serve as an inoculum reservoir for epidemics in wheat. In Brazil, severe wheat blast epidemics are most common in the Cerrado region. The dominant hypothesis is that signal grass (Urochloa sp.) and other gramineous plants harbor the wheat blast pathogen, thus serving as a major reservoir of inoculum for epidemics in wheat. A two-year survey of the Pyricularia blast pathogens was conducted in both wheat and non-wheat areas as well as prior (February) and during (May) the wheat growing season in Minas Gerais. A total of 1,368 plant samples representative of 31 Poaceae species, including wheat, were collected and inspected for the presence of blast symptoms. During the isolations, 932 isolates were obtained, being one fourth obtained from gramineous plants. A subset of 572 isolates was selected for identification at the species level based on portions of the CH7-BAC9 gene sequences. Most of the isolates (n = 494) were P. oryzae, within which 68% were PoT and 32% non-PoT based on two PCR assays targeting (MoT3 and C17 PCR assays). The PoT lineage was found predominantly (97%) in wheat and rarely in the other hosts, even nearby wheat fields (2.1%), as well as at longer distances from wheat regions (0.1%). The blast pathogen population isolated from signal grass grouped in different clades from PoT, and therefore referred to Urochloa lineage (PoU). A series of cross-inoculation greenhouse experiments was conducted using wheat (cv. BRS Guamirim and BR 18-Terena) and signal grass (cv. Marandu) as host and 14 PoT and six PoU isolates as pathogen factor. In the first leaf-inoculation experiment, results showed a significant interaction between host and pathogen; PoT was strongly/weakly aggressive towards wheat/signal grass and PoU was strongly/weakly aggressive towards signal grass/wheat. In inoculated wheat heads, PoT was more aggressive (>91% infected spikelets) than PoU (52% infected spikelets). In a third experiment, four signal grass cultivars (Marandu, Basilisk, Piatã, and Xaraés) were inoculated with the same set of 20 isolates. Similarly, signal grass cultivars were generally more susceptible to PoU than PoT. Severity induced by PoU was twice (7.7% severity) as high as PoT (3.8%) and so was the number of conidia/leaf produced by PoU (47,500) and PoT (23,200). Two groups of signal grass cultivars were formed, the most susceptible composed of Marandu and Basilisk and the least susceptible composed of Piatã and Xaraés. Results of our study confirm the host-specialization and the shaping of the blast populations according to the host. We further suggest that grasses in general, especially signal grass, may not play a major role as an inoculum reservoir for PoT, as it harbors mainly the PoU population. However, due to the large extent of pasture-growing regions and cross-infection ability in wheat, signal grass may harbor amounts of PoT inoculum that are sufficient for initiating leaf and head blast epidemics in wheat blast in Minas Gerais state.

GBS analysis of Orobanche crenata populations in Algeria supports local adaptation and host-specialization

Crenate broomrape (Orobanche crenata Forsk.) is a serious long-standing parasitic weed problem in Algeria, mainly affecting legumes but also vegetable crops. Unresolved questions for parasitic weeds revolve around the extent to which these plants undergo local adaptation, especially with respect to host specialization, which would be expected to be a strong selective factor for obligate parasitic plants. In the present study, the Genotyping-By-Sequencing (GBS) approach was used to analyze genetic diversity and population structure of 10 Algerian O. crenata populations with different geographical origins and host species (faba bean, pea, chickpea, carrot and tomato). In total, 8,004 high-quality single-nucleotide polymorphisms were obtained and used across the study. Genetic diversity and relationships of 95 individuals from 10 populations were studied using model-based ancestry analysis, principal components analysis, discriminant analysis of principal components, and phylogeny approaches. The genetic differentiation (FST) between pairs of populations was lower between adjacent populations and higher between geographically separated ones, but no support was found for isolation by distance. Further analyses identified four genetic clusters and revealed evidence of structuring among populations and hosts with more evident structuring among hosts than strictly along a geographic gradient. In the most striking example, O. crenata growing on pea had a distinct SNP profile from those growing on faba bean or other crops. These results illustrate the potential of GBS to reveal the dynamics of parasitic weed dispersal and adaptation.

Drosophila sechellia: A Genetic Model for Behavioral Evolution and Neuroecology

Defining the mechanisms by which animals adapt to their ecological niche is an important problem bridging evolution, genetics, and neurobiology. We review the establishment of a powerful genetic model for comparative behavioral analysis and neuroecology, Drosophila sechellia. This island-endemic fly species is closely related to several cosmopolitan generalists, including Drosophila melanogaster, but has evolved extreme specialism, feeding and reproducing exclusively on the noni fruit of the tropical shrub Morinda citrifolia. We first describe the development and use of genetic approaches to facilitate genotype/phenotype associations in these drosophilids. Next, we survey the behavioral, physiological, and morphological adaptations of D. sechellia throughout its life cycle and outline our current understanding of the genetic and cellular basis of these traits. Finally, we discuss the principles this knowledge begins to establish in the context of host specialization, speciation, and the neurobiology of behavioral evolution and consider open questions and challenges in the field. Expected final online publication date for the Annual Review of Genetics, Volume 55 is November 2021. Please see http://www.annualreviews.org/page/journal/pubdates for revised estimates.