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Key message: Multiple QTLs control unreduced pollen production in potato. Two major-effect QTLs co-locate with mutant alleles of genes with homology to AtJAS, a known regulator of meiotic spindle orientation. In diploid potato the production of unreduced gametes with a diploid (2n) rather than a haploid (n) number of chromosomes has been widely reported. Besides their evolutionary important role in sexual polyploidisation, unreduced gametes also have a practical value for potato breeding as a bridge between diploid and tetraploid germplasm. Although early articles argued for a monogenic recessive inheritance, the genetic basis of unreduced pollen production in potato has remained elusive. Here, three diploid full-sib populations were genotyped with an amplicon sequencing approach and phenotyped for unreduced pollen production across two growing seasons. We identified two minor-effect and three major-effect QTLs regulating this trait. The two QTLs with the largest effect displayed a recessive inheritance and an additive interaction. Both QTLs co-localised with genes encoding for putative AtJAS homologs, a key regulator of meiosis II spindle orientation in Arabidopsis thaliana. The function of these candidate genes is consistent with the cytological phenotype of mis-oriented metaphase II plates observed in the parental clones. The alleles associated with elevated levels of unreduced pollen showed deleterious mutation events: an exonic transposon insert causing a premature stop, and an amino acid change within a highly conserved domain. Taken together, our findings shed light on the natural variation underlying unreduced pollen production in potato and will facilitate interploidy breeding by enabling marker-assisted selection for this trait. [ABSTRACT FROM AUTHOR]

Aims: Zinc biofortification through agronomic and/or genetic approaches is one of the strategies to enrich food with this essential element to fulfil nutritional requirements in poor diets. This study aimed to unravel the physiological basis of differential tuber-Zn concentration between potato cultivars and to identify the major source(s) of Zn to the tubers. Methods: Two potato cultivars, accumulating high (Lady Rosetta) and low (Cara) tuber-Zn, were evaluated and the patterns of biomass and Zn accumulation between organs assessed over the growth period. Furthermore, Zn partition in reciprocal grafted plants and in sixteen F1 genotypes following a cross between those cultivars was assessed at harvest. Results: The cultivars differed in Zn-concentration, Zn-content and biomass partitioning between organs, although total plant-Zn content was comparable (4.7 and 5.1 mg Zn/plant for Cara and Lady Rosetta, respectively). At harvest the relative amount of Zn retained in roots, stems, leaves and tubers was 2 %,25 %,5 % and 68 % for Cara and 1 %,17 %,5 % and 77 % for Lady Rosetta. The differential tuber-Zn was associated with internal Zn distribution between shoots and tubers. Lady Rosetta translocated higher amounts of Zn to the tubers while Cara accumulated relatively more Zn in the stems associated with higher stem biomass. Remobilization of Zn from leaves and stems, and continuous root to xylem-phloem transfers during tuber bulking were the major sources of Zn supply to the tubers. Conclusions: This study suggests that the potato stem plays an important role in regulating Zn distribution and also as a site for Zn deposition. [ABSTRACT FROM AUTHOR]

Marker-assisted selection (MAS) is becoming increasingly common in commercial potato breeding programmes and can provide information to breeders on potential resistance to diseases and pests early in the breeding cycle when phenotypic screens are impractical. In recent years, a wealth of information on molecular markers for genes involved in resistance to biotic stresses of potato has been revealed. However, markers come in many different forms and ideally a breeding programme would routinely perform annual genotyping on a single unified platform. In this study, we present an approach to consolidate various types of molecular markers to a single SNP-based platform using whole-genome sequencing of carefully constructed pools of DNA using, as examples, molecular markers for the H1 nematode and R2 blight resistance genes. The approach led to the identification of a number of candidate nucleotide polymorphisms which allowed successful design and deployment of kompetitive allele-specific PCR (KASP) assays. We further show how the sequence data generated can be used to aid design of KASP assays for diagnostic SNPs for other traits mined from the literature. [ABSTRACT FROM AUTHOR]

Aim: Potato cultivars vary in biomass production and Cd concentration. Differences in growth can lead to differences in tuber cadmium (Cd) concentration through growth dilution and/or limited uptake of Cd from the soil. The aims of this study were to investigate the relative importance of shoot (above ground plant parts) and root in dry weight (DW) accumulation, total Cd uptake and Cd distribution between organs and how this affects tuber Cd concentration.Materials and methods: Three experiments were carried out, two with reciprocal grafting between high and low tuber Cd accumulating cultivars under two soil Cd conditions, and the other was involving the evaluation of F1 genotypes, from a cross between these high and low tuber Cd accumulating cultivars. These experiments were carried out under controlled greenhouse conditions.Results: The results showed that the root system is important for total Cd uptake but less important for growth and Cd distribution between organs. On the other hand, the shoot system is important for regulating growth of shoot and roots, and for determining Cd distribution between organs.Conclusion: The data suggest that tuber Cd concentration was mainly regulated by shoot source and to a lesser extent by total uptake of Cd. [ABSTRACT FROM AUTHOR]

Key message: Cd is a toxic metal, whilst Zn is an essential for plant and human health. Both can accumulate in potato tubers. We examine the genetic control of this process.Abstract: The aim of this study was to map quantitative trait loci (QTLs) influencing tuber concentrations of cadmium (Cd) and zinc (Zn). We developed a segregating population comprising 188 F1 progeny derived from crossing two tetraploid cultivars exhibiting divergent tuber-Cd-accumulation phenotypes. These progeny were genotyped using the SolCap 8303 SNP array, and evaluated for Cd, Zn, and maturity-related traits. Linkage and QTL mapping were performed using TetraploidSNPMap software, which incorporates all allele dosage information. The final genetic map comprised 3755 SNP markers with average marker density of 2.94 per cM. Tuber-Cd and Zn concentrations were measured in the segregating population over 2 years. QTL mapping identified four loci for tuber-Cd concentration on chromosomes 3, 5, 6, and 7, which explained genetic variance ranging from 5 to 33%, and five loci for tuber-Zn concentration on chromosome 1, 3, 5, and, 6 explaining from 5 to 38% of genetic variance. Among the QTL identified for tuber-Cd concentration, three loci coincided with tuber-Zn concentration. The largest effect QTL for both tuber-Cd and Zn concentration coincided with the maturity locus on chromosome 5 where earliness was associated with increased tuber concentration of both metals. Coincident minor-effect QTL for Cd and Zn sharing the same direction of effect was also found on chromosomes 3 and 6, and these were unrelated to maturity The results indicate partially overlapping genetic control of tuber-Cd and Zn concentration in the cross, involving both maturity-related and non-maturity-related mechanisms. [ABSTRACT FROM AUTHOR]

Potatoes grown in soil with high Cd concentrations can accumulate high levels of Cd in the tubers. Although there is significant environmental variation involved in the trait of crop uptake of Cd, there are also distinctive cultivar differences. In order to understand this differential Cd accumulation mechanism, two potato cultivars were chosen that accumulate high and low levels of Cd in tubers. The patterns of Cd concentration, Cd content and dry weight accumulation of the two cultivars were examined at different stages of plant growth. The data suggest that differences in total Cd uptake and in Cd partitioning among organs are the mechanisms governing differential Cd-tuber accumulation in the two cultivars. The low tuber-Cd accumulator exhibited lower root-to-shoot and shoot-to-tuber translocation driven by higher root and shoot biomass that retained more Cd in roots and shoots, respectively, reducing its movement to the tubers. Higher remobilization and more efficient tuber loading was observed in the high tuber-Cd accumulator, indicating that remobilization of Cd from leaves to tubers was a major factor, not only in tuber-Cd loading, but also in the establishment of differential tuber-Cd levels. Regardless of cultivar differences, the concentration of Cd in the tuber was very low compared to that in other organs suggesting that, despite its high phloem mobility, Cd tends to be sequestered in the shoots. [ABSTRACT FROM AUTHOR]

Globodera pallida is a major nematode pest causing severe constraints in many potato production regions worldwide. The most prevalent G. pallida pathotypes are Pa2 and Pa3, which exist in mixed populations referred to as Pa2/3. Due to heterogeneity for avirulence genes within these nematode populations, so far, breeders have failed to identify a single resistance gene source which offers complete resistance. Currently, there are two quantitative trait loci (QTL) available, $$ GpaIV_{adg}^{s} $$ and Gpa5, conferring partial levels of resistance to G. pallida pathotype Pa2/3. The objective of this research was to pyramid $$ GpaIV_{adg}^{s} $$ and Gpa5 through marker-assisted selection to investigate whether or not pyramiding provides increased resistance to G. pallida Pa2/3. We developed a population segregating for both resistance QTLs and, using diagnostic genetic markers (Contig237 and HC), we identified groups of individuals from this population containing each individual QTL, both QTLs simultaneously, and neither QTL. These individuals were assessed for their resistance levels against G. pallida Pa2/3 population Chavornay. We demonstrated that individuals carrying both QTLs showed a significant reduction in the number of cysts formed in comparison to genotypes carrying $$ GpaIV_{adg}^{s} $$ or Gpa5, indicating an additive effect. Overall these results show that MAS-based pyramiding of these QTLs is an effective strategy for breeding cultivars exhibiting very high levels of resistance to G. pallida pathotype Pa2/3. [ABSTRACT FROM AUTHOR]

White clover ( Trifolium repens L.) is a forage legume of considerable economic importance in temperate agricultural systems. It has a strong self-incompatibility system. The molecular basis of self-incompatibility in T. repens is unknown, but it is under the control of a single locus, which is expressed gametophytically. To locate the self-incompatibility locus ( S locus) in T. repens, we carried out cross-pollination experiments in an F1 mapping population and constructed a genetic linkage map using amplified fragment length polymorphism and simple sequence repeat markers. As the first step in a map-based cloning strategy, we locate for the first time the S locus in T. repens on a genetic linkage map, on the homoeologous linkage group pair 1 (E), which is broadly syntenic to Medicago truncatula L. chromosome 1. On the basis of this syntenic relationship, the possibility that the S locus may or may not possess an S- RNase gene is discussed. [ABSTRACT FROM AUTHOR]

Quantitative resistance to Globodera pallida pathotype Pa2/3, originally derived from Solanum tuberosum ssp. andigena Commonwealth Potato Collection (CPC) accession 2802, is present in several potato cultivars and advanced breeding lines. One genetic component of this resistance, a large effect quantitative trait locus (QTL) on linkage group IV (which we have renamed GpaIV) has previously been mapped in the tetraploid breeding line 12601ab1. In this study, we show that GpaIV is also present in a breeding line called C1992/31 via genetic mapping in an F1 population produced by crossing C1992/31 with the G. pallida susceptible cultivar Record. C1992/31 is relatively divergent from 12601ab1, confirming that GpaIV is an ideal target for marker-assisted selection in currently available germplasm. To generate markers exhibiting diagnostic potential for GpaIV , three bacterial artificial chromosome clones were isolated from the QTL region, sequenced, and used to develop 15 primer sets generating single-copy amplicons, which were examined for polymorphisms exhibiting linkage to GpaIV in C1992/31. Eight such polymorphisms were found. Subsequently, one insertion/deletion polymorphism, three single nucleotide polymorphisms and a specific allele of the microsatellite marker STM3016 were shown to exhibit diagnostic potential for the QTL in a panel of 37 potato genotypes, 12 with and 25 without accession CPC2082 in their pedigrees. STM3016 and one of the SNP polymorphisms, C237(119), were assayed in 178 potato genotypes, arising from crosses between C1992/31 and 16 G. pallida susceptible genotypes, undergoing selection in a commercial breeding programme. The results suggest that the diagnostic markers would most effectively be employed in MAS-based approaches to pyramid different resistance loci to develop cultivars exhibiting strong, durable resistance to G. pallida pathotype Pa2/3. [ABSTRACT FROM AUTHOR]

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Flowering time is a trait which has a major influence on the quality of forage. In addition, flowering and subsequent seed yields are important traits for seed production by grass breeders. In this study, we have identified quantitative trait loci (QTL) for flowering time and morphological traits of the flowering head in an F1 mapping population in Lolium perenne L (perennial ryegrass), a number of which have not previously been identified in L. perenne mapping studies. QTL for days to heading (DTH) were mapped in both outdoor and glasshouse experiments, revealing three and five QTL for DTH which explained 53% and 42% of the total phenotypic variation observed, respectively. Two QTL for DTH were detected in both environments, although they had contrasting relative magnitudes in each environment. One QTL for spike length and three QTL for spikelets per spike were also identified explaining, a total of 32 and 33% of the phenotypic variance, respectively. Furthermore, the QTL for spike length and spikelets per spike generally coincided with QTL for days to heading, implying co-ordinate regulation by underlying genes. Of particular interest was a region harbouring overlapping QTL for days to heading, spike length and spikelets per spike on the top of linkage group 4, containing the major QTL for spike length identified in this population. [ABSTRACT FROM AUTHOR]

Simple sequence repeat (SSR) markers are potentiallyvaluable tools for plant breeding. The objectives ofthe work reported here were to search the EMBL andGenbank databases for the presence of SSR-containingsequences from the genus Capsicum, to assess thefrequency of different motifs, and to examine thepolymorphism of selected markers in a panel ofgenotypes, including 10 Capsicum spp. and 1tomato and 1 potato genotype. Fifty-eightmicrosatellites with different motifs were found in Capsicum sequences. A subset of twelve of thesewere selected for the polymorphism survey using PCRprimers flanking the SSR. Polymorphisms between Capsicum lines can be detected with 5 of these primerpairs. PCR products of the predicted size were alsoamplified with three primer pairs in potato and oneprimer pair in tomato. The study also showed thatshorter microsatellites could be valuable markers in Capsicum. [ABSTRACT FROM AUTHOR]

The application of AFLPs, RAPDs and SSRs to examine genetic relationships in the primary northwestern European cultivated potato gene pool was investigated. Sixteen potato cultivars were genotyped using five AFLP primer combinations, 14 RAPD primers, and 17 database-derived SSR primer pairs. All three approaches successfully discriminated between the 16 cultivars using a minimum of one assay. Similarity matrices produced for each marker type on the basis of Nei and Li coefficients showed low correlations when compared with different statistical tests. Dendrograms were produced from these data for each marker system. The usefulness of each system was examined in terms of number of loci revealed (effective multiplex ratio, or EMR) and the amount of polymorphism detected (diversity index, or DI). AFLPs had the highest EMR, and SSRs the highest DI. A single parameter, marker index (MI), which is the product of DI and EMR, was used to evaluate the overall utility of each marker system. The use of these PCR-based marker systems in potato improvement and statutory applications is discussed. Abbreviations: PCR, polymerase chain reaction; AFLP, amplified fragment length polymorphism; RAPD, randomly amplified polymorphic DNA; DNA, deoxyribonucleic acid; EMR, effective multiplex ratio; DI, diversity index; MI, marker index; RFLP, restriction fragment length polymorphism. [ABSTRACT FROM AUTHOR]