Mapping and pyramiding of qualitative and quantitative resistance to stripe rust in barley

The identification and location of sources of genetic resistance to plant diseases are important contributions to the development of resistant varieties. The combination of different sources and types of resistance in the same genotype should assist in the development of durably resistant varieties. Using a doubled haploid (DH), mapping population of barley, we mapped a qualitative resistance gene (Rpsx) to barley stripe rust in the accession CI10587 (PI 243183) to the long arm of chromosome 1(7H). We combined the Rpsx gene, through a series of crosses, with three mapped and validated barley stripe rust resistance QTL alleles located on chromosomes 4(4H) (QTL4), 5(1H) (QTL5), and 7(5H) (QTL7). Three different barley DH populations were developed from these crosses, two combining Rpsx with QTL4 and QTL7, and the third combining Rpsx with QTL5. Disease severity testing in four environments and QTL mapping analyses confirmed the effects and locations of Rpsx, QTL4, and QTL5, thereby validating the original estimates of QTL location and effect. QTL alleles on chromosomes 4(4H) and 5(1H) were effective in decreasing disease severity in the absence of the resistance allele at Rpsx. Quantitative resistance effects were mainly additive, although magnitude interactions were detected. Our results indicate that combining qualitative and quantitative resistance in the same genotype is feasible. However, the durability of such resistance pyramids will require challenge from virulent isolates, which currently are not reported in North America.Communicated by J.W. SnapeOregon Agricultural Experiment Station paper No. 11953     

Molecular mapping of the Rph7.g leaf rust resistance gene in barley (Hordeum vulgare L.)

In many temperate areas of the world, leaf rust is becoming an important disease of barley. In the last decade, new races of Puccinia hordei G. Otth have emerged which are virulent against the so-far most-effective race-specific resistance genes, such as Rph7. Marker-assisted selection greatly facilitates the pyramidization of two or more resistance genes in a single variety in order to achieve a more comprehensive resistance. Such a strategy requires the development of efficient and reliable markers. Here, we have developed a linkage map and found RFLP markers closely linked to the Rph7.g resistance gene on chromosome 3HS of barley. The receptor-like kinase gene Hv3Lrk that maps at 3.2 cM from Rph7.g was used to develop a PCR-based marker by exploiting a single nucleotide polymorphism. This marker was detected in 11 out of 12 (92%) barley lines having Rph7 and represents a valuable tool for marker-assisted selection. In addition, the identification of markers flanking Rph7.g provides the basis for positional cloning of this gene. Received: 1 December 1999 / Accepted: 28 February 2000     

Identification and mapping of a new leaf stripe resistance gene in barley (Hordeum vulgare L.)

Pyrenophora graminea is the seed-borne pathogen causal agent of barley leaf stripe disease. Near-isogenic lines (NILs) carrying resistance of the cv ”Thibaut” against the highly virulent isolate Dg2 were obtained by introgressing the resistance into the genetic background of the susceptible cv ”Mirco”. The segregation of the resistance gene was followed in a F₂ population of 128 plants as well as on the F₃ lines derived from the F₂ plants; the segregation fitted the 1:2:1 ratio for a single gene. By using NILs, a RAPD marker associated with the resistance gene was identified; sequence-specific (STS) primers were designed on the basis of the amplicon sequence and a RILs mapping population with an AFLP-based map were used to position this molecular marker to barley chromosome 1 S (7HS). STS and CAPS markers were developed from RFLPs mapped to the telomeric region of barley chromosome 7HS and three polymorphic PCR-based markers were developed. The segregation of these markers was followed in the F₂ population and their map position with respect to the resistance gene was determined. Our results indicate that the Thibaut resistance gene, which we designated as Rdg2a, maps to the telomeric region of barley chromosome 7HS and is flanked by the markers OPQ-9₇₀₀ and MWG 2018 at distances of 3.1 and 2.5 cM respectively. The suitability of the PCR-based marker MWG2018 in selection- assisted barley breeding programs is discussed. Received: 22 June 2000 / Accepted: 16 October 2000     

QTL analysis of resistance to the rice blast pathogen in barley (Hordeum vulgare)

Barley is compatible with the rice blast pathogen (Pyricularia oryzae Cav.). Fiftyfour barley cultivars of diverse geographic origin and pedigree were inoculated with three isolates of the rice blast pathogen. All barley genotypes showed blast disease symptoms when inoculated at the seedling stage with each of the three isolates. However, one genotype showed quantitative resistance to all three isolates and three genotypes showed quantitative resistance to one or two of the isolates. By inoculating a set of doubled-haploid lines derived from the cross ’Harrington’ (susceptible) and ’TR306’ (resistant) with isolate Ken 54–20, we mapped quantitative trait loci (QTLs) determining seedling stage blast resistance. At all QTLs, TR306 contributed the resistance alleles. The four QTLs, when considered jointly, explained 43.6% of the phenotypic variation in blast symptom expression. A comparison of the blast resistance QTLs with other disease resistance QTLs reported in this population revealed a region on chromosome 4 (4H) with multiple disease resistance loci. It will be useful to capitalize on the syntenic relationship of rice and barley and to integrate information on species-specific resistance genes with information on the reaction of the two species to the same pathogen. Received: 7 January 2000 / Accepted: 22 September 2000     

Mapping of two genes for resistance to clubroot (Plasmodiophora brassicae) in a population of doubled haploid lines of Brassica oleracea by means of RFLP and AFLP markers

A genetic map covering 615 cM in 12 linkage groups was assembled based on 92 RFLP and AFLP markers segregating in a population of 107 doubled haploid lines (DH lines) of Brassica oleracea. The DH-line population was obtained through microspore culture from the of two homozygous parents: DH-line Bi derived from the cabbage landrace Bindsachsener, and DH-line Gr from broccoli cv ‘Greenia’. Sixty-five percent of the loci, and in some cases complete linkage groups, displayed distorted segregation ratios, a frequency much higher than that observed in populations of the same species. DH-line Bi was resistant to clubroot, which is caused by a Dutch field isolate of Plasmodiophora brassicae. Resistance in the DH-line population was determined in two ways: by assigning symptom grades to each plant, and by measuring the fresh weights of the healthy and affected parts of the root system of each plant. Using a multiple QTL mapping approach to analyze the fresh weight data, we found two loci for clubroot resistance; these were designated pb-3 and pb-4. The additive effects of these loci were responsible for 68% of the difference between the parents and for 60% of the genetic variance among DH-line means. Also, indications for the presence of two additional, minor QTLs were found. Analysis of symptom grades revealed the two QTLs pb-3 and pb-4, as well as one of the two minor QTLs indicated by analysis of the fresh weight data. Received: 29 April 1996 / Accepted: 10 May 1996     

Mapping of chromosome regions conferring boron toxicity tolerance in barley (Hordeum vulgare L.)

 Boron toxicity has been recognised as an important problem limiting production in the low-rainfall regions of southern Australia, West Asia and North Africa. Genetic variation for boron toxicity tolerance in barley has been characterised but the mode of inheritance and the location of genes controlling tolerance were not previously known. A population of 150 doubled-haploid lines from a cross between a boron toxicity tolerant Algerian landrace, Sahara 3771, and the intolerant Australian cultivar Clipper was screened in four tolerance assays. An RFLP linkage map of the Clipper×Sahara population was used to identify chromosomal regions associated with boron tolerance in barley. Interval regression-mapping allowed the detection of four chromosomal regions involved in the boron tolerance traits measured. A region on chromosome 2H was associated with leaf-symptom expression, a region on chromosome 3H was associated with a reduction of the affect of boron toxicity on root growth suppression, a region on chromosome 6H was associated with reduced boron uptake, and a region on chromosome 4H was also associated with the control of boron uptake as well as being associated with root-length response, dry matter production and symptom expression. The benefits and potential of marker-assisted selection for boron toxicity tolerance are discussed. Received: 18 December 1997 / Accepted: 28 November 1998     

Identification of QTLs for partial resistance to leaf rust (Puccinia hordei) in barley

 The partial resistance to leaf rust in barley is a quantitative resistance that is not based on hypersensitivity. To map the quantitative trait loci (QTLs) for partial resistance to leaf rust, we obtained 103 recombinant inbred lines (RILs) by single-seed descent from a cross between the susceptible parent L94 and the partially resistant parent Vada. These RILs were evaluated at the seedling and adult plant stages in the greenhouse for the latent period (LP) of the rust fungus, and in the field for the level of infection, measured as area under the disease progress curve (AUDPC). A dense genetic map based on 561 AFLP markers had been generated previously for this set of RILs. QTLs for partial resistance to leaf rust were mapped using the “Multiple Interval Mapping” method with the putative QTL markers as cofactors. Six QTLs for partial resistance were identified in this population. Three QTLs, Rphq1, Rphq2 and Rphq3, were effective at the seedling stage and contributed approximately 55% to the phenotypic variance. Five QTLs, Rph2, Rphq3, Rphq4, Rphq5, and/or Rphq6 contributed approximtely. 60% of the phenotypic variance and were effective at the adult plant stage. Therefore, only the QTLs Rphq2 and Rhpq3 were not plant-stage dependent. The identified QTLs showed mainly additive effects and only one significant interaction was detected, i.e. between Rphq1 and Rphq2. The map positions of these QTLs did not coincide with those of the race-specific resistance genes, suggesting that genes for partial resistance and genes for hypersensitive resistance represent entirely different gene families. Also, three QTLs for days to heading, of which two were also involved in plant height, were identified in the present recombinant inbred population. These QTLs had been mapped previously on the same positions in different populations. The perspectives of these results for breeding for durable resistance to leaf rust are discussed. Received: 15 July 1997 / Accepted: 30 December 1997     

Mapping genes for resistance to barley stripe rust (Puccinia striiformis f. sp. hordei)

Two genes conferring resistance to the barley stripe rust found in Mexico and South America, previously identified as race 24, were mapped to the M arms of barley chromosomes 7 and 4 in a doubled haploid population using molecular markers and the quantitative trait loci (QTL) mapping approach. The resistance gene on chromosome 7 had a major effect, accounting for 57% of the variation in disease severity. The resistance gene on chromosome 4 had a minor effect, accounting for 10% of the variation in trait expression. Two pairs of restriction fragment length polymorphism markers are being used to introgress the resistance genes to North American spring barley using molecular marker-assisted backcrossing.Ore. Agric Exp Stn J no. 10283     

RFLP mapping of BaYMV resistance gene rym3 in barley (Hordeum vulgare)

The rym3 (formerly designated ym3) gene conferring resistance to barley yellow mosaic virus (BaYMV) is effective against all strains of the virus but up to now has not been mapped to any chromosome. We performed a linkage analysis, using DNA extracted from individually harvested mature leaves of 153 F₂ plants derived from a cross between BaYMV-resistant cv ’Ishuku Shirazu’ carrying rym3 and susceptible cv ’Ko A’. Additionally, the F₃ lines derived from F₂ plants were grown in the BaYMV-infested field and examined for their reaction to BaYMV. Our results indicated that rym3 is located on the short arm of chromosome 5H and flanked by RFLP markers MWG28and ABG705A at distances of 7.2 and 11.7 cM, respectively. The chromosomal configuration estimated by DNA markers around rym3 and the utilization of these molecular markers for pyramiding with the BaYMV resistance genes in barley breeding programs are discussed. Received: 24 August 1998 / Accepted: 30 January 1999<@head-com-p1a.lf>Communicated by F. Salamini     

Several QTLs involved in osmotic-adjustment trait variation in barley (Hordeum vulgare L.)

 Osmotic adjustment (OA) was previously demonstrated to be an important adaptive mechanism of drought tolerance in cereals. In order to determine which genomic regions are involved in OA variation, 187 barley (Hordeum vulgare L.) recombinant inbred lines (RILs) derived from a cross between Tadmor (drought tolerant) and Er/Apm (susceptible) were studied in a growth chamber for their OA capacity (through correlated traits and by calculation), at an early growth stage and under two water treatments (soil moisture of 14% and 100% of field capacity). The continuous distribution of the traits and their broad-sense line heritabilities, ranging from 0.04 to 0.44, indicated that OA and related traits should have a polygenic nature. A subset of 167 RILs were also genotyped using 78 RFLP, 32 RAPD and three morphological markers and a linkage map was constructed. Despite strong environmental effects acting on the traits, interval mapping and single-marker ANOVA allowed the detection of three QTLs for relative water content (RWC), four QTLs for osmotic potential (ψ_π), two QTLs of osmotic potential at full turgor (ψ_π100) and one QTL for osmotic adjustment at a soil moisture of 14% field capacity. For the irrigated treatment, only two QTLs were detected: one for RWC and one for ψ_π100. Two chromosomal regions were involved in several OA-related trait variations and could be considered as regions controlling OA; these were present on chromosome 1 (7H) and chromosome 6 (6H), whereas other regions were specific for one trait. No major QTL was found. However, the genomic region involved in OA-related traits on chromosome 1 (7H) in barley seemed to be conserved for OA variation among cereals. Epistatic effects, with or without additive effects, acted on the traits. Received: 15 July 1997 / Accepted: 29 October 1997     

Mapping resistance to cereal aphids in barley

 A set of 150 doubled-haploid (DH) barley (Hordeum vulgare L.) lines derived from the cross of Harrington/TR306 was used to determine the number and chromosomal location of quantitative trait loci (QTLs) controlling resistance to cereal aphids. The experiments were conducted under natural infestation in the field during two growing seasons: 1994 and 1995. Aphid resistance was measured by counting the number of aphids per plot. Counts were made on a weekly basis. Each year at the time of maximum aphid density there was an obvious difference in reaction between the parental genotypes. The DH lines showed continuous variation for aphid density. Simple interval mapping and simplified composite interval mapping revealed that the principal QTL determining cereal aphid resistance is on the distal region of the short arm of chromosome 1. This aphid-resistance QTL is linked with a heading-date QTL. At the time of highest aphid infestation, this QTL accounted for 31% and 22% of the total variance of aphid density in 1994 and 1995, respectively. A QTL on chromosome 5 was also detected but only by simplified composite interval mapping. However, the largest consistent effect was due to the QTL on the short arm of chromosome 1. This QTL may be a useful target for marker-assisted selection for adult plant cereal aphid resistance in barley. Received: 10 September 1996/Accepted: 11 October 1996     

Localising QTLs for leaf rust resistance and agronomic traits in barley (Hordeum vulgare L.)

The Hordeum vulgare accession ’HOR 1063’ was crossed with the barley cultivar Krona, and 220 doubled haploid lines were produced based on this cross. A molecular map was constructed based on RFLP markers. Field trials were performed over 2 years and at two locations. In field trials, resistance to leaf rust by means of artificial infection, heading date, plant height and Kernel weight were assessed. For leaf rust resistance, 4 QTLs were localised, that explained 96.1% of the genetic variation. One QTL on chromosome 4H confirmed a position found in another genetic background and one mapped to the same position as Rph16 on chromosome 2H. All digenic effects decreased the effects of the respective QTLs. In addition to the denso-locus and the hex-v locus, other QTLs influencing heading date, plant length and kernel weight were found in this cross. Received: 16 July 1999 / Accepted: 7 September 1999     

Identification of polypeptide markers of barley yellow dwarf virus resistance and susceptibility genes in non-infected barley (Hordeum vulgare) plants

Summary Barley yellow dwarf (BYDV) is a group a closely related viruses which cause economic losses in a wide range of graminaceous species throughout the world. Barley plants can be protected from the effects of BYDV by the Yd2 resistance gene. Plants which contain the Yd2 gene also contain a constitutively expressed polypeptide which was not found in any plants without Yd2. Conversely, BYDV susceptible plants contain another constitutively expressed polypeptide which was not found in any of the BYDV-resistant lines examined. These two polypeptides appear to have the same molecular weight (as assessed by SDS-PAGE) and only slightly different iso-electric points. They also appear to contain an extensive range of similar antigenic determinants. Both polypeptides were found in F₁ hybrids made from resistant and susceptible plants. We suggest that these two polypeptides are the products of two allelic genes. Analysis of near-isogenic lines showed that the locus which controls the Yd2 resistance gene and the locus controlling the synthesis of the two polypeptides may be within ± 9 cM of each other. We have developed a Western blot technique which allows assessment of barley lines, 4-days after seed imbibition, for the presence of the Yd2 gene.     

A bacterial artificial chromosome library for barley (Hordeum vulgare L.) and the identification of clones containing putative resistance genes

Modern cultivated barley is an important cereal crop with an estimated genome size of 5000 Mb. To develop the resources for positional cloning and structural genomic analyses in barley, we constructed a bacterial artificial chromosome (BAC) library for the cultivar Morex using the cloning enzyme HindIII. The library contains 313344 clones (816 384-well plates). A random sampling of 504 clones indicated an average insert size of 106 kbp (range=30–195 kbp) and 3.4% empty vectors. Screening the colony filters for chloroplast DNA content indicated an exceptionally low 1.5% contamination with chloroplast DNA. Thus, the library provides 6.3 haploid genome equivalents allowing a >99% probability of recovering any specific sequence of interest. High-density filters were gridded robotically using a Genetix Q-BOT in a 4×4 double-spotted array on 22.5-cm² filters. Each set of 17 filters allows the entire library to be screened with 18432 clones represented per filter. Screening the library with 40 single copy probes identified an average 6.4 clones per probe, with a range of 1–13 clones per probe. A set of resistance-gene analog (RGA) sequences identified 121 RGA-containing BAC clones representing 20 different regions of the genome with an average of 6.1 clones per locus. Additional screening of the library with a P-loop disease resistance primer probe identified 459 positive BAC clones. These data indicate that this library is a valuable resource for structural genomic applications in barley. Received: 20 September 1999 / Accepted: 25 March 2000     

QTL analysis of tolerance to a German strain of BYDV-PAV in barley (Hordeum vulgare L.)

One hundred and forty six barley doubled-haploid lines (DH lines) were tested for variation in grain yield, yield components, plant height, and heading date after artificial infection with a German isolate of barley yellow dwarf virus (BYDV-PAV-Braunschweig). Of these 146 lines 76 were derived from the cross of the barley yellow dwarf virus (BYDV) tolerant cultivar ’Post’ to cv ’Vixen’ (Ryd2) and 70 from the cross of Post to cv ’Nixe’. Phenotypic measurements were gathered on both non-infected plants and plants artificially inoculated with BYDV-PAV by viruliferous aphids in pot and field experiments for three years at two locations. For all traits a continuous variation was observed suggesting a quantitative mode of inheritance for tolerance against BYDV-PAV. Using skeleton maps constructed using SSRs, AFLPs and RAPDs, two QTLs for relative grain yield per plant after BYDV infection, explaining about 47% of the phenotypic variance, were identified in Post × Vixen at the telomeric region of chromosome 2HL and at a region containing the Ryd2 gene on chromosome 3HL. In Post × Nixe, a QTL was found in exactly the same chromosome 2HL marker interval. In this cross, additional QTL were mapped on chromosomes 7H and 4H and together these explained about 40% of the phenotypic variance. QTL for effects of BYDV infection on yield components, plant height, and heading date generally mapped to the same marker intervals, or in the vicinity of the QTL for relative grain yield, on chromosomes 2HL and 3HL, suggesting that these regions are of special importance for tolerance to the Braunschweig isolate of BYDV-PAV. Possible applications of marker-assisted selection for BYDV tolerance based on these results are discussed. Received: 1 December 2000 / Accepted: 9 March 2001     

Optimization of gene transfer into barley (Hordeum vulgare L.) mature embryos by tissue electroporation

 This study was conducted to detect the optimum conditions for DNA transfer into mature embryos of barley via electroporation. Cultured mature embryos of barley were directly electroporated in the presence of the pBI 121 vector carrying both the β-glucuronidase (GUS) and neomycin phosphotransferase II (npt II) genes. It was found that 500 v/cm and 500 μFd capacitance was the optimum combination for healthy germination of the transformed plants from mature electroporated embryos. Effects of culture duration before electroporation and selection antibiotic concentrations on germination were also examined. Gene transfer performed on 3-day-old cultures resulted in the highest germination frequencies. GUS expression was observed on transversal sections of embryos and mature leaves from 3 month-old regenerants. PCR and Southern blot analyses show the presence of the npt II transgene in the genome of a plant. Received: 15 June 1999 / Revision received: 27 September 1999 / Accepted: 26 October 1999     

Histological analysis of somatic embryogenesis in Brazilian cultivars of barley, Hordeum vulgare vulgare, Poaceae

 Histological analysis was performed aimed at elucidating the origin and the developmental process of somatic embryos of two Brazilian cultivars of barley (Hordeum vulgare vulgare), 'MN-599' and 'A-05'. The observed site of somatic embryo origin (SSEO) could originate from a superficial callus cell, possibly indicating a unicellular origin, or from epidermal and subepidermal callus cells, representing a multicellular origin. A fold, the somatic embryo scutellum that subsequently develops into a cotyledonary leaf, indicates the somatic embryo differentiation. The somatic embryos also showed a growth increase of the primary root and, occasionally, a delay in root development. A possible alternative pathway for the origin of somatic embryos is suggested, in which a SSEO forms a clump of somatic embryos. Received: 4 June 1998 / Revision received: 28 August 1998 / Accepted: 7 December 1998     

Polymorphism, distribution, and segregation of AFLP markers in a doubled haploid rice population

We exploited the newly developed amplified fragment length polymorphism (AFLP) technique to study the polymorphism, distribution and inheritance of AFLP markers with a doubled haploid rice population derived from ‘IR64’/‘Azucena’. Using only 20 pairs of primer combinations, we detected 945 AFLP bands of which 208 were polymorphic. All 208 AFLP markers were mapped and distributed over all 12 chromosomes. When these were compared with RFLP markers already mapped in the population, we found the AFLP markers to be highly polymorphic in rice and to follow Mendelian segregation. As linkage map of rice can be generated rapidly with AFLP markers they will be very useful for marker-assisted backcrossing. Received: 11 April 1996 / Accepted: 14 June 1996     

A molecular,isozyme and morphological map of the barley (Hordeum vulgare) genome

A map of the barley genome consisting of 295 loci was constructed. These loci include 152 cDNA restriction fragment length polymorphism (RFLP), 114 genomic DNA RFLP, 14 random amplified polymorphic DNA (RAPD), five isozyme, two morphological, one disease resistance and seven specific amplicon polymorphism (SAP) markers. The RFLP-identified loci include 63 that were detected using cloned known function genes as probes. The map covers 1,250 centiMorgans (cM) with a 4.2 cM average distance between markers. The genetic lengths of the chromosomes range from 124 to 223 cM and are in approximate agreement with their physical lengths. The centromeres were localized to within a few markers on all of the barley chromosomes except chromosome 5. Telomeric regions were mapped for the short (plus) arms of chromosomes 1, 2 and 3 and the long (minus) arm of chromosomes 7.This research was also supported by other members of the NABGMP: K. Kasha, Department of Crop Science, University of Guelph, Guelph, Ontario, Canada NIG 2W1; W. Kim, Agriculture Canada Research Station, 195 Dafoe Road, Winnipeg, Manitoba, Canada R3T 2M9; A. Laroche, Agriculture Canada Research Station, P.O. Box 3000 Main, Lethbridge, Alberta, Canada,TU 4B1; S. Molnar, Plant Research Centre Agriculture Canada, Central Experimental farm, Ottawa, Ontario, Canada K1A 0C6; G. Scoles, Department of Crop Science, University of Saskatchewan, Saskatoon, Saskatchewan, Canada S7N OWOThis research is part of the North American Barley Genome Mapping Project, R. A. Nilan and K. Kasha, Coordinator and Associate Coordinator, respectively Permanent address: Department of Plant Genetics, NI Vavilov Institute of General Genetics, Russian Academy of Sciences, Moscow     

Identification of two key genes controlling chill haze stability of beer in barley (Hordeum vulgare L)

Background

In bright beer, haze formation is a serious quality problem, degrading beer quality and reducing its shelf life. The quality of barley (Hordeum vulgare L) malt, as the main raw material for beer brewing, largely affects the colloidal stability of beer.

Results

In this study, the genetic mechanism of the factors affecting beer haze stability in barley was studied. Quantitative trait loci (QTL) analysis of alcohol chill haze (ACH) in beer was carried out using a Franklin/Yerong double haploid (DH) population. One QTL, named as qACH, was detected for ACH, and it was located on the position of about 108 cM in chromosome 4H and can explain about 20 % of the phenotypic variation. Two key haze active proteins, BATI-CMb and BATI-CMd were identified by proteomics analysis. Bioinformatics analysis showed that BATI-CMb and BATI-CMd had the same position as qACH in the chromosome. It may be deduced that BATI-CMb and BATI-CMd are candidate genes for qACH, controlling colloidal stability of beer. Polymorphism comparison between Yerong and Franklin in the nucleotide and amino acid sequence of BATI-CMb and BATI-CMd detected the corresponding gene specific markers, which could be used in marker-assisted selection for malt barley breeding.

Conclusions

We identified a novel QTL, qACH controlling chill haze of beer, and two key haze active proteins, BATI-CMb and BATI-CMd. And further analysis showed that BATI-CMb and BATI-CMd might be the candidate genes associated with beer chill haze.

Electronic supplementary material

The online version of this article (doi:10.1186/s12864-015-1683-1) contains supplementary material, which is available to authorized users.