Bochra A. Bahri

Genetic structure and local adaptation of European wheat yellow rust populations: the role of temperature-specific adaptation

Environmental heterogeneity influences coevolution and local adaptation in host–parasite systems. This also concerns applied issues, because the geographic range of parasites may depend on their capacity to adapt to abiotic conditions. We studied temperature‐specific adaptation in the wheat yellow/stripe rust pathogen, Puccinia striiformis f.sp. tritici (PST). Using laboratory experiments, PST isolates from northern and southern France were studied for their ability to germinate and to infect bread and durum wheat cultivars over a temperature gradient. Pathogen origin × temperature interactions for infectivity and germination rate suggest local adaptation to high‐ versus low‐temperature regimes in south and north. Competition experiments in southern and northern field sites showed a general competitive advantage of southern over northern isolates. This advantage was particularly pronounced in the southern ‘home’ site, consistent with a model integrating laboratory infectivity and field temperature variation. The stable PST population structure in France likely reflects adaptation to ecological and genetic factors: persistence of southern PST may be due to adaptation to the warmer Mediterranean climate; and persistence of northern PST can be explained by adaptation to commonly used cultivars, for which southern isolates are lacking the relevant virulence genes. Thus, understanding the role of temperature‐specific adaptations may help to improve forecast models or breeding programmes.

Tracking costs of virulence in natural populations of the wheat pathogen, Puccinia striiformis f.sp.tritici.

BackgroundCosts of adaptation play an important role in host-parasite coevolution. For parasites, evolving the ability to circumvent host resistance may trade off with subsequent growth or transmission. Such costs of virulence (sensu plant pathology) limit the spread of all-infectious genotypes and thus facilitate the maintenance of genetic polymorphism in both host and parasite. We investigated costs of three virulence factors in Puccinia striiformis f.sp.tritici, a fungal pathogen of wheat (Triticum aestivum).ResultsIn pairwise competition experiments, we compared the fitness of near-isogenic genotypes that differed by a single virulence factor. Two virulence factors (vir4, vir6) imposed substantial fitness costs in the absence of the corresponding resistance genes. In contrast, the vir9 virulence factor conferred a strong competitive advantage to several isolates, and this for different host cultivars and growing seasons. In part, the experimentally derived fitness costs and benefits are consistent with frequency changes of these virulence factors in the French pathogen population.ConclusionOur results illustrate the variation in the evolutionary trajectories of virulence mutations and the potential role of compensatory mutations. Anticipation of such variable evolutionary outcomes represents a major challenge for plant breeding strategies. More generally, we believe that agro-patho-systems can provide valuable insight in (co)evolutionary processes in host-parasite systems.

Geographic limits of a clonal population of wheat yellow rust in the Mediterranean region

Most plant pathogens present complex life cycles, in which the clonal reproduction may impede the delimitation of population entities. By studying the genetic structure of the wheat yellow rust caused by Puccinia striiformis f.sp. tritici (PST), we highlighted difficulties impeding population delimitation in highly clonal species. Despite the high dispersal potential of PST, southern France isolates were shown to be divergent from a northwestern European population. A 2‐year survey was performed in the Mediterranean region to assess the geographic distribution of southern isolates: 453 isolates collected in 11 countries were genotyped using 15 simple sequence repeat markers. A subsample was analysed for virulence against 23 resistance genes. The dominant strain in the western Mediterranean region was further studied with amplified fragment length polymorphism markers to test for a geographic substructure. Both ‘individual’‐ and ‘population’‐centred analyses of polymorphism markers revealed two south‐specific groups: a predominant group, with a broad variability and a wide distribution in both western and eastern Mediterranean countries, and a minor group in the western Mediterranean. The east–west gradient of genetic diversity suggested gene flow from the Middle East with subsequent founder effects and genetic divergence, and demonstrated the local survival of a western Mediterranean population. The high frequency of the resistance gene Yr8 observed in cultivars from Tunisia and Algeria may contribute to maintain the north/south structure observed in France. In addition to migration and local adaptation, the dynamics of clonal lineage diversification and replacement should be considered to define population entities in strongly clonal species.

Population genetics of Setaria viridis, a new model system

An extensive survey of the standing genetic variation in natural populations is among the priority steps in developing a species into a model system. In recent years, green foxtail (Setaria viridis), along with its domesticated form foxtail millet (S. italica), has rapidly become a promising new model system for C4 grasses and bioenergy crops, due to its rapid life cycle, large amount of seed production and small diploid genome, among other characters. However, remarkably little is known about the genetic diversity in natural populations of this species. In this study, we survey the genetic diversity of a worldwide sample of more than 200 S. viridis accessions, using the genotyping‐by‐sequencing technique. Two distinct genetic groups in S. viridis and a third group resembling S. italica were identified, with considerable admixture among the three groups. We find the genetic variation of North American S. viridis correlates with both geography and climate and is representative of the total genetic diversity in this species. This pattern may reflect several introduction/dispersal events of S. viridis into North America. We also modelled demographic history and show signal of recent population decline in one subgroup. Finally, we show linkage disequilibrium decay is rapid (<45 kb) in our total sample and slow in genetic subgroups. These results together provide an in‐depth understanding of the pattern of genetic diversity of this new model species on a broad geographic scale. They also provide key guidelines for on‐going and future work including germplasm preservation, local adaptation, crossing designs and genomewide association studies.

Isolation of ten microsatellite loci in an EST library of the phytopathogenic fungus Puccinia striiformis f.sp. tritici

We report the characterization of ten microsatellite markers in the fungus Puccinia striiformis f.sp. tritici, responsible for yellow rust disease on wheat. A published EST library was scanned for microsatellite motives, and over 15 selected EST sequences, 13 were successfully amplified and ten exhibited polymorphism over an international collection of 43 isolates. These new microsatellites, added to the few previously published ones, provide a sufficient set of markers to perform population genetic studies.

Molecular markers for tracking the origin and worldwide distribution of invasive strains of Puccinia striiformis

Abstract Investigating the origin and dispersal pathways is instrumental to mitigate threats and economic and environmental consequences of invasive crop pathogens. In the case of Puccinia striiformis causing yellow rust on wheat, a number of economically important invasions have been reported, e.g., the spreading of two aggressive and high temperature adapted strains to three continents since 2000. The combination of sequence‐characterized amplified region (SCAR) markers, which were developed from two specific AFLP fragments, differentiated the two invasive strains, PstS1 and PstS2 from all other P. striiformis strains investigated at a worldwide level. The application of the SCAR markers on 566 isolates showed that PstS1 was present in East Africa in the early 1980s and then detected in the Americas in 2000 and in Australia in 2002. PstS2 which evolved from PstS1 became widespread in the Middle East and Central Asia. In 2000, PstS2 was detected in Europe, where it never became prevalent. Additional SSR genotyping and virulence phenotyping revealed 10 and six variants, respectively, within PstS1 and PstS2, demonstrating the evolutionary potential of the pathogen. Overall, the results suggested East Africa as the most plausible origin of the two invasive strains. The SCAR markers developed in the present study provide a rapid, inexpensive, and efficient tool to track the distribution of P. striiformis invasive strains, PstS1 and PstS2.

Genetic diversity and origin of North American green foxtail [Setaria viridis (L.) Beauv.] accessions

AbstractSetaria viridis (L.) P. Beauv. and its domesticated form, S. italica (L.) P. Beauv., have been developed over the past few years as model systems for C4 photosynthesis and for the analysis of bioenergy traits. S. viridis is native to Eurasia, but is now a ubiquitous weed. An analysis of the population structure of a set of 232 S. viridis lines, mostly from North America but also comprising some accessions from around the world, using 11 SSR markers, showed that S.viridis populations in the US largely separate by latitude and/or climatic zone. S. viridis populations from the Northern US and Canada (north of 44°N) group with accessions from Western Europe, while populations in the Mid and Southern US predominantly group with accessions from Turkey and Iran. We hypothesize that S. viridis in the US was most likely introduced from Europe, and that introductions were competitive only in regions that had climatic conditions that were similar to those in the regions of origins. This hypothesis is supported by the fact that Canadian S. viridis lines were fast cycling and undersized when grown in the Mid-Western and Southern US compared to their morphology in their native environment. A comparison of the population structure obtained with 11 SSR markers and ~40,000 single nucleotide polymorphisms (SNPs) in a common set of S. viridis germplasm showed that both methods essentially yielded the same groupings, although admixture was identified at a higher frequency in the SNP analysis. Small numbers of SSR markers can thus be used effectively to discern the population structure in this inbreeding species.

Identification of resistance sources to Septoria tritici blotch in old Tunisian durum wheat germplasm applied for the analysis of the Zymoseptoria tritici-durum wheat interaction.

Septoria tritici blotch (STB) caused by the fungus Zymoseptoria tritici (formerly known as Mycosphaerella graminicola) is currently the most important foliar disease of durum wheat (Triticum turgidum subsp. durum) in Tunisia, causing serious yield losses and affecting grain quality. Resistance breeding in durum wheat to STB can provide an effective, economic and environmentally-safe strategy to reduce yield losses. However, this is hampered by a lack of resistance sources, and a limited understanding of Z. tritici pathogenicity. Here, we report the identification of nine resistant accessions to STB upon field inoculation of 144 old local durum wheat accessions with a virulent isolate “TunBz-1”. Crosses between the resistant accessions and evaluation of F2 progenies for their reaction to TunBz-1 led to the identification of four new resistant genes, associated with Azizi27, Agili37, Agili39 and Derbessi12 landrace accessions. Random distribution of the resistant accessions in an AFLP based-dendrogram of 123 old durum wheat accessions suggested independent resistance gene-evolution. Based on the identified resistance sources, the analysis of the virulence spectrum of 55 Z. tritici isolates collected from different durum wheat- growing regions in Tunisia showed a highly significant genotype-by-isolate interaction. Furthermore, pathotype variability among Z. tritici isolates was low as the isolates were classified into two groups (I and II) that showed differential reaction on the accession Azizi27. Our results show that the old local durum wheat germplasm might harbor novel resistance genes that can be deployed in durum wheat breeding programs.

Thermal generalist behaviour of invasive Puccinia striiformis f. sp. tritici strains under current and future climate conditions

Yellow rust is a devastating wheat disease. Since 2000, Puccinia striiformis f. sp. tritici strains PstS1 and PstS2 have become adapted to high temperatures and have spread worldwide. By 2011, Warrior strains had invaded both warm and cold areas of Europe. This study questioned whether thermal aptitude promoted the spread of Warrior strains, similar to PstS1/PstS2, by comparing infection efficiency (IE) at five temperatures and latent period (LP) under warm and cold regimes for Warrior isolates and pre‐2011 reference strains on two susceptible wheat varieties. The Warrior isolates showed a range of IE and LP responses to temperature that was intermediate between the northern reference isolates adapted to cold conditions and both the southern and invasive PstS2 isolates adapted to warm conditions. Warrior isolates had the highest IE under optimal temperatures of 10 and 15 °C, and displayed reduced infectivity under the warmest (20 °C) and coldest (5 °C) temperatures. Warrior strains acted as thermal generalists and the reference isolates acted as specialists. An IE thermal response was used to simulate the development of each isolate under future climate scenarios in a temperate and Mediterranean region. Isolates had the same ranking for yearly IE over the three 30‐year periods (1971–2000, 2021–2050, 2071–2100) and both locations, with a slight infection increase in the future. However, in the future IEs increased in earlier months. The thermal generalist profile of Warrior isolates for IE was confirmed, with an intermediate capacity to tolerate warming climate, whereas the southern isolates are better adapted to warm conditions, but do not have the virulences necessary to develop on current varieties.

QTL mapping combined with comparative analyses identified candidate genes for reduced shattering in Setaria italica

Setaria (L.) P. Beauv is a genus of grasses that belongs to the Poaceae (grass) family, subfamily Panicoideae. Two members of the Setaria genus, Setaria italica (foxtail millet) and S. viridis (green foxtail), have been studied extensively over the past few years as model species for C4-photosynthesis and to facilitate genome studies in complex Panicoid bioenergy grasses. We exploited the available genetic and genomic resources for S. italica and its wild progenitor, S. viridis, to study the genetic basis of seed shattering. Reduced shattering is a key trait that underwent positive selection during domestication. Phenotyping of F2:3 and recombinant inbred line (RIL) populations generated from a cross between S. italica accession B100 and S. viridis accession A10 identified the presence of additive main effect quantitative trait loci (QTL) on chromosomes V and IX. As expected, enhanced seed shattering was contributed by the wild S. viridis. Comparative analyses pinpointed Sh1 and qSH1, two shattering genes previously identified in sorghum and rice, as potentially underlying the QTL on Setaria chromosomes IX and V, respectively. The Sh1 allele in S. italica was shown to carry a PIF/Harbinger MITE in exon 2, which gave rise to an alternatively spliced transcript that lacked exon 2. This MITE was universally present in S. italica accessions around the world and absent from the S. viridis germplasm tested, strongly suggesting a single origin of foxtail millet domestication. The qSH1 gene carried two MITEs in the 5′UTR. Presence of one or both MITEs was strongly associated with cultivated germplasm. If the MITE insertion(s) in qSH1 played a role in reducing shattering in S. italica accessions, selection for the variants likely occurred after the domestication of foxtail millet.