杨树抗锈性研究现状

综述了落叶松一杨栅锈菌生理小种分化、寄主杨树抗锈性、抗锈病基因分子标记和抗病基因连锁遗传图谱构建技术;分析了我国杨树抗锈病育种研究的现状和亟待解决的问题。     

Inheritance of leaf rust and stem rust resistance in 'Roblin' wheat.

The Canadian common wheat (Triticum aestivum L.) cultivar 'Roblin' is resistant to both leaf rust (Puccinia recondita Rob. ex. Desm.) and stem rust (Puccinia graminis Pers. f. sp. tritici Eriks. and E. Henn.). To study the genetics of this resistance, 'Roblin' was crossed with 'Thatcher', a leaf rust susceptible cultivar, and RL6071, a stem rust susceptible line. A set of F6 random lines was developed from each cross. The random lines and the parents were grown in a field rust nursery artificially inoculated with a mixture of P. recondita and P. graminis isolates and scored for rust reaction. The same material was tested with specific races of leaf rust and stem rust. These data indicated that 'Roblin' has Lr1, Lr10, Lr13, and Lr34 for resistance to P. recondita and Sr5, Sr9b, Sr11, and possibly Sr7a and Sr12 for resistance to P. graminis. In a 'Thatcher' background, the presence of Lr34 contributes to improve stem rust resistance, which appears also to occur in 'Roblin'.     

A novel gene for rust resistance

The Rpg1 gene, which has provided North American cultivars of barley with resistance to the stem rust fungus Puccinia graminis f.sp. tritici for more than 60 years, has been cloned. A single copy of the gene can confer resistance to a susceptible barley variety. Although unexplained, the progeny are consistently more resistant than the variety from which the gene was obtained. The gene might represent a new class of plant resistance genes.     

Nonhost resistance of rice to rust pathogens

Rice is atypical in that it is an agricultural cereal that is immune to fungal rust diseases. This report demonstrates that several cereal rust species (Puccinia graminis f. sp tritici, P. triticina, P. striiformis, and P. hordei) can infect rice and produce all the infection structures necessary for plant colonization, including specialized feeding cells (haustoria). Some rust infection sites are remarkably large and many plant cells are colonized, suggesting that nutrient uptake occurs to support this growth. Rice responds with an active, nonhost resistance (NHR) response that prevents fungal sporulation and that involves callose deposition, production of reactive oxygen species, and, occasionally, cell death. Genetic variation for the efficacy of NHR to wheat stem rust and wheat leaf rust was observed. Unlike cereal rusts, the rust pathogen (Melampsora lini) of the dicotyledenous plant flax (Linum usitatissimum) rarely successfully infects rice due to an apparent inability to recognize host-derived signals. Morphologically abnormal infection structures are produced and appressorial-like structures often don't coincide with stomata. These data suggest that basic compatibility is an important determinate of nonhost infection outcomes of rust diseases on cereals, with cereal rusts being more capable of infecting a cereal nonhost species compared with rust species that are adapted for dicot hosts.     

Autoactive alleles of the flax L6 rust resistance gene induce non-race-specific rust resistance associated with the hypersensitive response

L6 is a nucleotide binding site-leucine rich repeat (NBS-LRR) gene that confers race-specific resistance in flax (Linum usitatissimum) to strains of flax rust (Melampsora lini) that carry avirulence alleles of the AvrL567 gene but not to rust strains that carry only the virulence allele. Several mutant and recombinant forms of L6 were made that altered either the methionine-histidine-aspartate (MHD) motif conserved in the NBS domain of resistance proteins or exchanged the short domain C-terminal to the LRR region that is highly variable among L allele products. In transgenic flax some of these alleles are autoactive; they cause a gene dosage-dependent dwarf phenotype and constitutive expression of genes that are markers for the plant defense response. Their effects and penetrance ranged from extreme to mild in their degree of plant stunting, survival, and reproduction. Dwarf plants were also resistant to flax rust strains virulent to wild-type L6 plants, and this nonspecific resistance was associated with a hypersensitive response (HR) at the site of rust infection. The strongest autoactive allele, expressed in Arabidopsis from an ethanol-inducible promoter, gave rise to plant death dependent on the enhanced disease susceptibility 1 (EDS1) gene, which indicates that the mutant flax (Linaceae) L6 gene can signal cell death through a defined disease-resistance pathway in a different plant family (Brassicaceae).     

Durable resistance to wheat stem rust needed

The recent outbreak of a new wheat stem rust race capable of parasitizing many commercial wheat cultivars highlights the need for durable disease resistance in crop plants. More advanced breeding approaches using quantitative disease resistance genes and resistance gene pyramids are being used to combat wheat stem rust and other diseases, though widespread adoption of these breeding methodologies is needed to maintain resistance efficacy. Advances in understanding the molecular basis of plant disease resistance at both host and nonhost levels offers further possibilities for stem rust resistance using biotechnological approaches. However, truly durable resistance to wheat stem rust and other phytopathogens seems an unlikely prospect in the face of continually evolving pathogen populations.     

Stripe rust and leaf rust resistance QTL mapping,epistatic interactions,and co-localization with stem rust resistance loci in spring wheat evaluated over three continents

Key message

In wheat, advantageous gene-rich or pleiotropic regions for stripe, leaf, and stem rust and epistatic interactions between rust resistance loci should be accounted for in plant breeding strategies.

Abstract

Leaf rust (Puccinia triticina Eriks.) and stripe rust (Puccinia striiformis f. tritici Eriks) contribute to major production losses in many regions worldwide. The objectives of this research were to identify and study epistatic interactions of quantitative trait loci (QTL) for stripe and leaf rust resistance in a doubled haploid (DH) population derived from the cross of Canadian wheat cultivars, AC Cadillac and Carberry. The relationship of leaf and stripe rust resistance QTL that co-located with stem rust resistance QTL previously mapped in this population was also investigated. The Carberry/AC Cadillac population was genotyped with DArT^® and simple sequence repeat markers. The parents and population were phenotyped for stripe rust severity and infection response in field rust nurseries in Kenya (Njoro), Canada (Swift Current), and New Zealand (Lincoln); and for leaf rust severity and infection response in field nurseries in Canada (Swift Current) and New Zealand (Lincoln). AC Cadillac was a source of stripe rust resistance QTL on chromosomes 2A, 2B, 3A, 3B, 5B, and 7B; and Carberry was a source of resistance on chromosomes 2B, 4B, and 7A. AC Cadillac contributed QTL for resistance to leaf rust on chromosome 2A and Carberry contributed QTL on chromosomes 2B and 4B. Stripe rust resistance QTL co-localized with previously reported stem rust resistance QTL on 2B, 3B, and 7B, while leaf rust resistance QTL co-localized with 4B stem rust resistance QTL. Several epistatic interactions were identified both for stripe and leaf rust resistance QTL. We have identified useful combinations of genetic loci with main and epistatic effects. Multiple disease resistance regions identified on chromosomes 2A, 2B, 3B, 4B, 5B, and 7B are prime candidates for further investigation and validation of their broad resistance.     

Gene-for-gene-mediated resistance to southern corn rust in maize

Southern corn rust (SCR) severely threatens maize production worldwide. Achieving durable control of SCR requires efficient breeding and deployment of resistant hybrids. Recently, two research teams (Chen et al. and Deng et al.) cloned two SCR resistance genes (RppC and RppK) and the cognate Avr genes (AvrRppC and AvrRppK), which will accelerate SCR resistance breeding.     

Homologues of the maize rust resistance gene Rp1-D are genetically associated with a major rust resistance QTL in sorghum

As part of a comparative mapping study between sugarcane and sorghum, a sugarcane cDNA clone with homology to the maize Rp1-D rust resistance gene was mapped in sorghum. The cDNA probe hybridised to multiple loci, including one on sorghum linkage group (LG) E in a region where a major rust resistance QTL had been previously mapped. Partial sorghum Rp1-D homologues were isolated from genomic DNA of rust-resistant and -susceptible progeny selected from a sorghum mapping population. Sequencing of the Rp1-D homologues revealed five discrete sequence classes: three from resistant progeny and two from susceptible progeny. PCR primers specific to each sequence class were used to amplify products from the progeny and confirmed that the five sequence classes mapped to the same locus on LG E. Cluster analysis of these sorghum sequences and available sugarcane, maize and sorghum Rp1-D homologue sequences showed that the maize Rp1-D sequence and the partial sugarcane Rp1-D homologue were clustered with one of the sorghum resistant progeny sequence classes, while previously published sorghum Rp1-D homologue sequences clustered with the susceptible progeny sequence classes. Full-length sequence information was obtained for one member of a resistant progeny sequence class ( Rp1-SO) and compared with the maize Rp1-D sequence and a previously identified sorghum Rp1 homologue ( Rph1-2). There was considerable similarity between the two sorghum sequences and less similarity between the sorghum and maize sequences. These results suggest a conservation of function and gene sequence homology at the Rp1 loci of maize and sorghum and provide a basis for convenient PCR-based screening tools for putative rust resistance alleles in sorghum.     

High-density genetic mapping for coffee leaf rust resistance

Coffee leaf rust caused by the fungus Hemileia vastatrix causes considerable economic losses for coffee producers. Although agrochemical products can provide sufficient disease control, the use of resistant cultivars is a safer alternative. This resistance may be constrained by one or a few genetic factors, mainly those found in material originating from interspecific hybrids. In this study, the genetic analysis of an F ₂ population consisting of 224 plants derived from a crossing of Híbrido de Timor UFV 427-15 (resistant) with Catuaí Amarelo IAC 30 (susceptible) showed that a dominant gene confers the resistance of coffee to race II of H. vastatrix. From a genetic map saturated with 25 amplified fragment length polymorphism (AFLP) markers linked to the resistance gene, we developed a high-density genetic map with six sequence-characterized amplified region (SCAR) markers delimiting a chromosomal region of 9.45 cM and flanking the dominant gene at 0.7 and 0.9 cM. This is the first saturated and high-density genetic map obtained from this region containing the resistance gene. The results of this study are of great importance for the introduction of molecular markers for marker-assisted selection; they will also facilitate studies related to the cloning, structure, and function of race-specific genes involved in the resistance of coffee trees to H. vastatrix.     

QTL mapping of resistance to leaf rust in Salix

Genetic mapping of quantitative trait loci (QTL) for resistance to Melampsora leaf rust was performed in two willow families: the progeny from a backcross between Salix viminalis and a hybrid S. viminalis × Salix schwerinii (population S₁), and the F₁ progeny of a cross between two S. viminalis (population S₃). Disease levels were scored in the field for three consecutive years. In the laboratory, five different rust strains were sprayed onto leaf disks and the following resistance components were scored: latent period, diameter and number of uredinia, and flecking. One major QTL and 14 smaller were identified in the S₁ host population. One rust strain, that represents a Melampsora form with limited incidence on S. viminalis, showed lower aggressiveness to the S₁ host population together with a different pattern in QTLs. In the S₃ host population, we detected 13 QTLs for rust resistance, of which two were located to the same genomic regions as those found for the S₁ population. We showed that the strongest QTL co-segregated with a gene homologous to a known Toll/interleukin receptor-nucleotide binding site-leucine-rich repeat resistance gene in poplar. The importance of the identified QTLs is discussed in relation to breeding for durable resistance.     

Genetic mapping of rust resistance loci in biomass willow

Rust diseases caused by Melampsora spp. represent a major threat to the productivity of short rotation coppice (SRC) willows grown for biomass, causing yield losses of up to 40%. The routine use of fungicide in SRC plantations is not a viable option because of economic and environmental considerations; thus, breeding for rust resistance is a major target for willow breeding programmes. To characterise the genetic basis of rust resistance in willow and provide targets for use in future marker-assisted selections, quantitative trait analyses were performed using a large full-sib mapping population (K8) which segregates for rust resistance and several other important agronomic traits. Rust resistance in field conditions was assessed in three consecutive years. For a more detailed genetic dissection, laboratory inoculation tests using isolates of two distinct and prevalent pathotypes (LET1 and LET5) were also performed. For field-based resistance, a major quantitative resistance locus, designated SRR1 (Salix Rust Resistance 1), was detected in addition to several quantitative trait loci (QTL) of more modest effect. Inoculation test data also supported an important role for SRR1. Specific interactions between particular rust isolates and different QTL were detected, and QTL that only influenced resistance in field conditions were identified. The QTL reported here represent an important basis for the future development of markers for use in willow breeding programmes. As the linkage map for the K8 population is anchored to the Populus trichocarpa genome sequence, a more efficient marker development for future fine-scale mapping and candidate gene identification is possible.     

Quantitative trait loci of stripe rust resistance in wheat

Key message

Over 140 QTLs for resistance to stripe rust in wheat have been published and through mapping flanking markers on consensus maps, 49 chromosomal regions are identified.

Abstract

Over thirty publications during the last 10 years have identified more than 140 QTLs for stripe rust resistance in wheat. It is likely that many of these QTLs are identical genes that have been spread through plant breeding into diverse backgrounds through phenotypic selection under stripe rust epidemics. Allelism testing can be used to differentiate genes in similar locations but in different genetic backgrounds; however, this is problematic for QTL studies where multiple loci segregate from any one parent. This review utilizes consensus maps to illustrate important genomic regions that have had effects against stripe rust in wheat, and although this methodology cannot distinguish alleles from closely linked genes, it does highlight the extent of genetic diversity for this trait and identifies the most valuable loci and the parents possessing them for utilization in breeding programs. With the advent of cheaper, high throughput genotyping technologies, it is envisioned that there will be many more publications in the near future describing ever more QTLs. This review sets the scene for the coming influx of data and will quickly enable researchers to identify new loci in their given populations.     

Leaf rust resistance genes of wheat: identification in cultivars and resistance sources

Thirty-seven wheat cultivars originating from seven European countries were examined by using sequence tagged site (STS) markers for seven Lr (leaf rust = brown rust) resistance genes against the fungal pathogen of wheat Puccinia recondita f. sp. tritici (Lr9, Lr10, Lr19, Lr24, Lr26 and Lr37). Additionally, 22 accessions with various Lr genes from two germplasm collections were tested. A Scar (sequence-characterized amplified region) marker for Lr24 and a CAPS (Cleaved Amplified Polymorphic Sequence) marker for Lr47 were also used to identify those genes in the wheat accessions. Each marker amplified one specific DNA fragment. Three Lr gene markers were identified in wheat cultivars (Lr10, Lr26 and Lr37). Another four markers (Lr9, Lr19, Lr24 and Lr47) were found in breeding lines carrying leaf rust resistance genes. The results were compared with leaf rust resistance gene postulations made in previous studies, based on multipathotype testing. Markers for Lr10, Lr26 and Lr37 may be useful in marker-assisted breeding.     

Mapping of quantitative adult plant field resistance to leaf rust and stripe rust in two European winter wheat populations reveals co-location of three QTL conferring resistance to both rust pathogens

Key message

We detected several, most likely novel QTL for adult plant resistance to rusts. Notably three QTL improved resistance to leaf rust and stripe rust simultaneously indicating broad spectrum resistance QTL.

Abstract

The rusts of wheat (Puccinia spp.) are destructive fungal wheat diseases. The deployment of resistant cultivars plays a central role in integrated rust disease management. Durability of resistance would be preferred, but is difficult to analyse. The Austrian winter wheat cultivar Capo was released in the 1989 and grown on a large acreage during more than two decades and maintained a good level of quantitative leaf rust and stripe rust resistance. Two bi-parental mapping populations: Capo × Arina and Capo × Furore were tested in multiple environments for severity of leaf rust and stripe rust at the adult plant stage in replicated field experiments. Quantitative trait loci associated with leaf rust and stripe rust severity were mapped using DArT and SSR markers. Five QTL were detected in multiple environments associated with resistance to leaf rust designated as QLr.ifa-2AL, QLr.ifa-2BL, QLr.ifa-2BS, QLr.ifa-3BS, and QLr.ifa-5BL, and five for resistance to stripe rust QYr.ifa-2AL, QYr.ifa-2BL, QYr.ifa-3AS, QYr.ifa-3BS, and QYr.ifa-5A. For all QTL apart from two (QYr.ifa-3AS, QLr.ifa-5BL) Capo contributed the resistance improving allele. The leaf rust and stripe rust resistance QTL on 2AL, 2BL and 3BS mapped to the same chromosome positions, indicating either closely linked genes or pleiotropic gene action. These three multiple disease resistance QTL (QLr.ifa-2AL/QYr.ifa-2AL, QLr.ifa.2BL/QYr.ifa-2BL, QLr.ifa-3BS/QYr.ifa.3BS) potentially contribute novel resistance sources for stripe rust and leaf rust. The long-lasting resistance of Capo apparently rests upon a combination of several genes. The described germplasm, QTL and markers are applicable for simultaneous resistance improvement against leaf rust and stripe rust.     

Genetics of durable resistance to leaf rust and stripe rust of an Indian wheat cultivar HD2009

The Indian bread wheat cultivar HD2009 has maintained its partial resistance to leaf rust and stripe rust in India since its release in 1976. To examine the nature, number and mode of inheritance of its genes for partial leaf rust and stripe rust resistance, this cultivar was crossed with cultivar WL711, which is susceptible to leaf rust and stripe rust. The F1, F2, F3 and F5 generations from this cross were assessed separately for adult plant disease severity under artificial epidemic of race 77-5 of leaf rust and race 46S119 of stripe rust. Segregation for rust reaction in the F2, F3 and F5 generations indicated that resistance to each of these rust diseases is based on 2 genes, each with additive effects. Although the leaf rust resistance of HD2009 is similar in expression to that conferred by the gene Lr34, but unlike the wheats carrying this gene, cultivar HD2009 did not show leaf tip necrosis, a morphological marker believed to be tightly linked to the leaf rust resistance gene Lr34. Thus, the non-hypersensitive resistance of HD2009 was ascribed to genes other than Lr34.     

Breeding for mature-plant resistance to yellow rust in wheat

In recent years several varieties of wheat, such as Rothwell Perdix and Maris Envoy, have shown good resistance to yellow rust when first produced, but because their resistance was determined by simply inherited major genes, they have later been severely attacked by newly arising physiologic races of the pathogen. Other varieties, such as Little Joss, Atle and Maris Widgeon, though slightly attacked by several races, have never suffered severe damage even when exposed to high levels of inoculum. The inheritance of this non-race-specific type of resistance was studied in a cross between Little Joss and a susceptible but shorter-strawed variety, Nord Desprez. Resistance appeared to be under complex control. It was found possible to select short-strawed resistant plants, using simple techniques in the field.     

Identification of the stem rust resistance gene Pg9 and its association with crown rust resistance and endosperm proteins in 'Dumont' oat.

The Canadian oat cultivar 'Dumont' is known to have genes Pc38 and Pc39 for crown rust resistance and genes Pg2 and Pg13 for stem rust resistance. When crossed to a susceptible oat line OT328, 'Dumont' was shown to have an additional dominant gene for crown rust resistance, designated PcX. Tests of segregating progeny indicated that the stem rust resistance gene Pg9 is present and is tightly linked in coupling to PcX. The presence of Pg9 in 'Dumont' was confirmed in crosses involving the cultivar 'Ukraine', which has Pg9 and a crown rust resistance gene tightly linked to it. The association of rust resistance with endosperm proteins in 'Dumont' was investigated. The linkage of gene Pg13 with a 56.6-kDa polypeptide locus (map distance of 10.47 +/- 2.70 cM) was demonstrated using sodium dodecylsulfate - polyacrylamide gel electrophoresis (SDS-PAGE). A 27.9-kDa polypeptide was shown to be associated with the linked PcX/Pg9 loci by SDS-PAGE but appeared to be more reliably separated as an avenin band, designated B4, using acid-PAGE. Another avenin band, designated B2, also was shown to be associated with the PcX/Pg9 loci using acid-PAGE. The loci conditioning the B2 and B4 bands appeared to be tightly linked or allelic and are separated from the linked PcX/Pg9 loci by a map distance of 1.03 +/- 0.36 cM. The association of Pg13 with a 56.6-kDa polypeptide and the tight linkage between PcX/Pg9 and the B2 (in coupling) and B4 (in repulsion) avenin loci offer a useful tool to breeders to detect the presence of these genes in oat breeding.     

QTL characterization of resistance to leaf rust and stripe rust in the spring wheat line Francolin#1

Growing resistant wheat varieties is a key method of controlling two important wheat diseases, leaf rust and stripe rust. We analyzed quantitative trait loci (QTL) to investigate adult plant resistance (APR) to these rusts, using 141 F₅ RILs derived from the cross ‘Avocet-YrA/Francolin#1’. Phenotyping of leaf rust resistance was conducted during two seasons at Ciudad Obregon, Mexico, whereas stripe rust was evaluated for two seasons in Toluca, Mexico, and one season in Chengdu, China. The genetic map was constructed with 581 markers, including diversity arrays technology and simple sequence repeat. Significant loci for reducing leaf rust severity were designated QLr.cim-1BL, QLr.cim-3BS.1, QLr.cim-3DC, and QLr.cim-7DS. The six QTL that reduced stripe rust severity were designated QYr.cim-1BL, QYr.cim-2BS, QYr.cim-2DS, QYr.cim-3BS.2, QYr.cim-5AL, and QYr.cim-6AL. All loci were conferred by Francolin#1, with the exception of QYr.cim-2DS, QYr.cim-5AL, and QYr.cim-6AL, which were derived from Avocet-YrA. Closely linked markers indicated that the 1BL locus was the pleiotropic APR gene Lr46/Yr29. QYr.cim-2BS was a seedling resistance gene designated as YrF that conferred intermediate seedling reactions and moderate resistance at the adult plant stage in both Mexican and Chinese environments. Significant additive interactions were detected between the six QTL for stripe rust, but not between the four QTL for leaf rust. Furthermore, we detected two new APR loci for leaf rust in common wheat: QLr.cim-3BS.1 and QLr.cim-7DS.