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SummaryIn the Netherlands, sweet pepper (Capsicum annuum L.) plants are grown in heated greenhouses. Lowering the greenhouse temperature results in a substantial reduction in energy use. In order to grow crops under lowered temperatures, cultivars are required which are adapted to such conditions. In this study, we compared the variance in adaptation to low temperatures in six cultivated sweet pepper varieties to that in a collection of ten wild and cultivated Capsicum accessions. The adaptation to lowered temperatures was defined in terms of variation in the difference in relative growth rate (RGR) between plants grown under different temperature conditions. Our results illustrated that within the group of sweet peppers, the variation in low-temperature tolerance was limited, and that sweet pepper breeders should exploit the genetic variation in wild relative species in order to breed for this trait. RGR, however, is difficult and expensive to assess. Therefore, we also studied which morphological plant ...

In plants, several population types [F(2), recombinant inbred lines, backcross inbred lines (BILs), etc.] are used for quantitative trait locus (QTL) analyses. However, dissection of the trait of interest and subsequent confirmation by introgression of QTLs for breeding purposes has not been as successful as that predicted from theoretical calculations. More practical knowledge of different QTL mapping approaches is needed. In this recent study, we describe the detection and mapping of quantitative resistances to downy mildew in a set of 29 BILs of cultivated lettuce (L. sativa) containing genome segments introgressed from wild lettuce (L. saligna). Introgression regions that are associated with quantitative resistance are considered to harbor a QTL. Furthermore, we compare this with results from an already existing F(2) population derived from the same parents. We identified six QTLs in our BIL approach compared to only three in the F(2) approach, while there were two QTLs in common. We performed a simulation study based on our actual data to help us interpret them. This revealed that two newly detected QTLs in the BILs had gone unnoticed in the F(2), due to a combination of recessiveness of the trait and skewed segregation, causing a deficit of the wild species alleles. This study clearly illustrates the added value of extended genetic studies on two different population types (BILs and F(2)) to dissect complex genetic traits.

Use of DNA-based markers can accelerate cultivar development in variable cultivation environments since, in contrast to phenotype, DNA markers are environment-independent. In an effort to elucidate the genetic basis of genotype-by-environment interaction (G x E) for yield of rice (Oryza sativa L.), the associations between 139 AFLP markers and grain yield were determined for rice grown in fresh water (EC of 0.65 dS m(-1) and saline conditions (EC of 4-8 dS m(-1)) with 0 kg ha(-1) or 100 kg ha(-1) nitrogen fertilizer in the years 2000 and 2001. A population of recombinant inbred lines of rice, developed from an IR29 x Pokkali cross, was used in the study. Both genotype x salinity and genotype x nitrogen level interactions were significant, with the genotype x salinity interaction being stronger. Through multiple regression analysis using a stepwise procedure for selecting markers, 36 markers were detected for grain yield in the four test conditions and of these 28 were detected in only one test condition implying strong environmental specificity for yield QTL expression. However, the fact that eight QTLs were detected in more than one test condition points to the existence of wide-adaptability genes in this cross. Markers with significant associations with yield explained between 37% and 48% of the yield variation in each test condition. Superior genotypes of rice were identified in all four test conditions based on their marker signatures. Furthermore, across N fertilizer regimes, yield predicted from summed additive effects of QTLs were significantly correlated with observed yield in the same year and across years. Thus marker-assisted selection can help breeders overcome the problem of low selection efficiency encountered during phenotypic selection for yield in stress environments.

During meiosis, homologous chromosomes (homologs) perform reciprocal exchanges (crossovers) at a high frequency. Crossovers display interference, i.e. their spacing is more even than would be expected if they were placed randomly along the chromosomes. Concomitantly with crossover formation, synaptonemal complexes (SCs) appear between homologs: each chromosome forms an axial structure, the axial element (AE); the AEs of homologs align, and numerous transverse filaments connect the AEs to form an SC. Both the AE and the SC have been implicated in the imposition of interference. We investigated whether intact AEs or SCs are required for crossover interference in the mouse, using a mutant lacking AE protein SYCP3, which displays structurally abnormal AEs and incomplete synapsis. We estimated the level of interference from the spacing of immunofluorescent MLH1 foci, which mark almost all crossover sites in the mouse, along the SCs. The levels of interference among MLH1 foci in wild-type and Sycp3–/– mice were comparable, implying that neither an intact AE structure nor full synapsis is required for wild-type levels of interference.

The improvement of energy efficiency in the greenhouse production of cut rose and pot rose can be achieved through the use of rose cultivars having vigorous growth. A better understanding of the inheritance of vigour and its related traits will assist the breeding activities. Quantitative trait locus (QTL) analyses were performed with the help of an integrated linkage map of a diploid rose population originating from a cross between Rosa multiflora-derived genotypes. The underlying datasets for ten vigour-related traits were collected in an evaluation study of this population in two greenhouse experiments with suboptimal temperatures for growth. We identified ten chromosomal regions, scattered over the seven linkage groups, containing QTLs for these traits. Considering each trait separately, we detected a total of 42 QTLs. Among these QTLs, 24 were found in both of the experiments, eight and ten were specific to either of the two experiments. The number of QTLs for individual traits varied from three to five with a respective contribution to the phenotypic variation from 12 to 35%. QTLs for highly correlated traits frequently co-localized, indicating a common genetic basis. Clustering of QTLs for different traits was noted in some chromosome regions, for instance, one on chromosome 2 included major QTLs for eight of ten traits under study, suggesting co-localization of several separate genes or/and the occurrence of various genes having pleiotropic effects. The discovery of markers associated to QTL regions is in roses the first step towards marker-assisted selection for vigour improvement enabling the transfer of useful QTL-alleles of R. multiflora to pot and cut roses.

A quantitative trait locus analysis was carried out to unravel the genetic basis of dormancy in wild barley (Hordeum spontaneum) from Israel. Two accessions, Ashkelon and Mehola, from divergent environments were crossed to produce a mapping population. A linkage map was produced from the F2 population, and F4 seeds were used for germination experiments. Five quantitative trait loci (QTL) were detected for dormancy across the different germination experiments. These QTL were found on chromosomes 1, 2, 5, 6 and 7. The variation explained by each QTL varied between 8 and 25%. Ashkelon alleles increased the germination except for the QTL on chromosome 5. Three out of these five QTL co-locate with QTL found earlier in cultivated barley crosses, although this does not necessarily imply that they would be the same loci. The level of dormancy is much higher in wild barley than in cultivated barley and wild barley may have alleles that have not yet been utilised in breeding for optimally dormant barley.

Powdery mildew (Podosphaera pannosa) is the most important fungal disease in greenhouse roses and is in practice controlled by fungicides. The creation of novel cultivars with durable resistance to powdery mildew is highly desirable. To understand the inheritance of mildew resistance, a tetraploid rose population with a size of 181 seedlings was obtained by crossing two tetraploid genotypes each having partial resistance. The population and its parents were tested under greenhouse conditions with two well-defined monospore isolates (2 and F1) using artificial inoculation with spore suspensions. Disease score at 11 days post-inoculation, latent period and rate of symptom development were used to describe seedling resistance. The tests for both isolates exhibited a wide and significant variation among genotypes for resistance. The distribution of the genotypic means of the disease scores was continuous and showed a considerable transgression. Statistical analysis, scatter plot of disease scores for the isolates, and correlation analyses indicated that the two isolates differed in pathogenicity. The outcome of the tests showed that the inoculation assay with spore suspensions was a reliable and effective way to screen large numbers of genotypes under greenhouse conditions for genetic and breeding studies. This is the first report on spore-suspension inoculation to be used successfully in rose.

During meiosis, homologous chromosomes (homologs) undergo recombinational interactions, which can yield crossovers (COs) or noncrossovers. COs exhibit interference; they are more evenly spaced along the chromosomes than would be expected if they were placed randomly. The protein complexes involved in recombination can be visualized as immunofluorescent foci. We have analyzed the distribution of such foci along meiotic prophase chromosomes of the mouse to find out when interference is imposed and whether interference manifests itself at a constant level during meiosis. We observed strong interference among MLH1 foci, which mark CO positions in pachytene. Additionally, we detected substantial interference well before this point, in late zygotene, among MSH4 foci, and similarly, among replication protein A (RPA) foci. MSH4 foci and RPA foci both mark interhomolog recombinational interactions, most of which do not yield COs in the mouse. Furthermore, this zygotene interference did not depend on SYCP1, which is a transverse filament protein of mouse synaptonemal complexes. Interference is thus not specific to COs but may occur in other situations in which the spatial distribution of events has to be controlled. Differences between the distributions of MSH4/RPA foci and MLH1 foci along synaptonemal complexes might suggest that CO interference occurs in two successive steps.

A droplet inoculation method was used for evaluation of tomato resistance to early blight, a destructive foliar disease of tomato caused by Alternaria solani (Ellis and Martin) Sorauer. In this test method, leaflets are inoculated with small droplets of a spore suspension in either water or a 0.1% agar solution. Early blight resistance was evaluated based on lesion size. The droplet method better discriminated the level of resistance (P < 0.001) for a range of spore densities in comparison with the more commonly used spray inoculation method. Lesions generated by droplet inoculation at 7 days after inoculation ranged from small flecks to almost complete blight with an exponential-like distribution of lesion sizes. Significant correlations (r = 0.52, 0.58, and 0.63, P < 0.001) were observed across three glasshouse tests of 54 accessions including wild species using the droplet method. The most resistant accessions included wild species: one accession of Solanum arcanum, three accessions of Solanum peruvianum, one accession of Solanum neorickii, and one of Solanum chilense. Solanum pennellii and Solanum pimpinellifolium accessions were susceptible, whereas Solanum habrochaites and Solanum lycopersicum accessions ranged from susceptible to moderately resistant. The droplet test method is simple to apply, offers a fine discrimination of early blight resistance levels, and allows objective evaluation.

Plant growth can be studied at different organizational levels, varying from cell, leaf, and shoot to the whole plant. The early growth of seedlings is important for the plant's establishment and its eventual success. Wheat (Triticum aestivum, genome AABBDD) seedlings exhibit a low early growth rate or early vigor. The germplasm of wheat is limited. Wild relatives constitute a source of genetic variation. We explored the physiological and genetic relationships among a range of early vigor traits in Aegilops tauschii, the D-genome donor. A genetic map was constructed with amplified fragment-length polymorphism and simple sequence repeat markers, and quantitative trait loci (QTL) analysis was performed on the F4 population of recombinant inbred lines derived from a cross between contrasting accessions. The genetic map consisted of 10 linkage groups, which were assigned to the seven chromosomes and covered 68% of the D genome. QTL analysis revealed 87 mapped QTLs (log of the odds >2.65) in clusters, 3.1 QTLs per trait, explaining 32% of the phenotypic variance. Chromosomes 1D, 4D, and 7D harbored QTLs for relative growth rate, biomass allocation, specific leaf area, leaf area ratio, and unit leaf rate. Chromosome 2D covered QTLs for rate and duration of leaf elongation, cell production rate, and cell length. Chromosome 5D harbored QTLs for the total leaf mass and area and growth rate of the number of leaves and tillers. The results show that several physiological correlations between growth traits have a genetic basis. Genetic links between traits are not absolute, opening perspectives for identification of favorable alleles in A. tauschii to improve early vigor in wheat.

Accurate measurements of leaf area are important for agronomic and physiological studies. To be able to perform repeated measurements of leaf area on single (genetically unique) plants, a method wa...

Parental linkage maps of a segregating population of diploid rose hybrids (2n=2x =14), composed of 365 uni-parental AFLP and SSR markers, have been constructed using a population (n=88) derived from a cross between two half-sib parents (P119 and P117). Of the markers, 157 P119 markers (85 %) mapped on eight linkage groups and 133 P117 markers (78 %) on seven linkage groups. The resulting linkage maps of P119 and P117 spanned 463 cM and 491 cM with an average of interval between markers of 2.9 cM and 3.7 cM, respectively. The present genetic maps were used to identify quantitative trait loci (QTLs) for two growth vigour-related traits, leaf area and chlorophyll content, using the Multiple QTL Mapping approach. Three QTLs for leaf area and two QTLs for chlorophyll content were identified. The QTLs accounted, in total, for 50.8 % (range 7.0-23.1 %) and 25.8 % (range 7.6-18.2 %) of the total phenotypic variance for leaf area and chlorophyll content, respectively. The detection of highly significant major QTLs enables marker-assisted selection for growth vigour in rose.

A method is presented for the selection of parents with the aim of obtaining improved genotypes in the progeny of a cross. The procedure is designed to select in several unrelated traits simultaneously and is based on the selection of molecular markers that are linked to QTLs. The method was compared with conventional phenotypic selection in simulation experiments for a number of genetic structures underlying the traits and several types of parental populations. Although the method in general provides good results, some of the underlying assumptions may be violated quite easily, thereby reducing the applicability of the procedure in practice.

Using AFLP markers, a linkage map was constructed based on a recombinant inbred population of barley derived from a cross between a leaf rust susceptible line, L94, and a partially resistant line, 116-5. The constructed map showed a similar marker distribution pattern as the L94 × Vada map. However, it contained more large gaps, and for some chromosome regions no markers were identified. These regions are most likely derived from L94 because 116-5 was selected from the progeny of a cross of L94 × cv. Cebada Capa. Five QTLs for partial resistance to isolate 1.2.1. were mapped on the L94 × 116-5 map. Three QTLs were effective in the seedling stage, jointly contributing 42% to the total phenotypic variance. Three QTLs were effective in the adult plant stage, collectively explaining 35% of the phenotypic variance. Evidence for two additional linked minor-effect QTLs effective in the adult plant stage was also uncovered. The major-effect QTL, Rphq3, was the only one that was effective in both developmental stages. Moreover, Rphq3, was also identified in the L94 × Vada population, being effective to two rust isolates. The other QTLs were detected in either of the two populations, providing evidence for the existence of many loci for partial resistance to leaf rust on the barley genome. To date, 13 QTLs for partial resistance have been mapped, therefore, a strategy of accumulating many resistance genes in a single cultivar, resulting in a high level of partial resistance, is feasible.

An amplified fragment length polymorphism (AFLP) map covering 965 cM was constructed using 94 recombinant inbred lines of a cross between the spring barley varieties Prisma and Apex. This map was employed to identify quantitative trait loci (QTLs) controlling plant height, yield and yield-determining physiological characters using an approximate multiple-QTL model, the MQM method. The seven physiological traits were parameters used in a process-based crop-growth model that predicts barley biomass production as affected by daily temperature and radiation. The traits were measured in experiments conducted over 2 years. Except for the relative growth rate of leaf area, all traits examined had at least one QTL in each year. QTLs and their effects were found to vary with developmental stages for one trait, the fraction of shoot biomass partitioned to leaves, that was measured at several stages. Most of the traits were associated, though to different extents, with the denso dwarfing gene (the height-reducing allele in Prisma) located on the long arm of chromosome 3. Some of the QTLs were mapped to similar positions in both years. The results in relation to effects of the dwarfing gene, the physiological basis for QTL x environment interaction, and the relative importance of the parameter traits with respect to yield, are discussed.

Crop modelling has so far contributed little to the genetic analysis of a quantitative trait. This study illustrates how a simple model for crop phenological development, which assumes that crop development rate is affected by daily effective temperature, can assist the identification of Quantitative Trait Loci (QTLs), using specific leaf area (SLA) in barley as an example. The SLA was measured in a field experiment six times during the growing season of 94 recombinant inbred lines (RILs) derived from a cross between cultivars Prisma and Apex. Of the six measurements, one was conducted at the same physiological age for all RILs (at flowering), four were undertaken at specific chronological days prior to flowering, and the last one was taken at 14 days after flowering. When the measured SLA was directly used as the quantitative trait, one to three QTLs were detected for SLA at each measurement time. The major dwarfing gene denso segregating in the population was found to affect SLA strongly at all measurement times except at flowering. If SLA of the different RILs was corrected for differences in physiological age at the time of measurement, by the use of the crop development model, QTLs were detected for SLA at only three stages. Furthermore, the effect of the denso gene was no longer significant during the preflowering stages. The effect of the denso gene detected in the first instance was therefore the consequence of its direct effect on the duration of the preflowering period. This demonstrates the important role that crop development models can play in QTL analysis of a trait that varies with developmental stage. Potential uses of ecophysiological crop growth models in QTL analysis are briefly discussed.

Molecular markers may enable plant breeders to select indirectly for genes affecting quantitative traits by selecting for molecular markers closely linked to these genes (marker-assisted selection, MAS). We have assessed the effectiveness of MAS compared to phenotypic selection. Key variables in this assessment were: trait heritability, selection intensity, genetic architecture and uncertainty in QTL mapping. Simulation studies showed that the application of MAS in autogamous crops, with the objective of obtaining transgressive genotypes, can improve selection results when compared to conventional selection procedures. Marker-assisted selection appears particularly promising when dominant alleles at quantitative trait loci are present and linked in coupling phase. Uncertainty in estimated QTL map positions reduces the benefits of marker-assisted selection, but this reduction remains limited in most cases.

By using 25 primer combinations, 563 AFLP markers segregating in a recombinant inbred population (103 lines, F9) derived from L94/Vada were generated. The 38 AFLP markers in common to the existing AFLP/RFLP combined Proctor/Nudinka map, one STS marker, and four phenotypic markers with known map positions, were used to assign present AFLP linkage groups to barley chromosomes. The constructed high-density molecular map contains 561 AFLP markers, three morphological markers, one disease resistance gene and one STS marker, and covers a 1062-cM genetic distance, corresponding to an average of one marker per 1.9 cM. However, extremely uneven distributions of AFLP markers and strong clustering of markers around the centromere were identified in the present AFLP map. Around the centromeric region, 289 markers cover a genetic distance of 155 cM, corresponding to one marker per 0.5 cM; on the distal parts, 906 cM were covered by 277 markers, corresponding to one marker per 3.3 cM. Three gaps larger than 20 cM still exist on chromosomes 1, 3 and 5. A skeletal map with a uniform distribution of markers can be extracted from the high-density map, and can be applied to detect and map loci underlying quantitative traits. However, the application of this map is restricted to barley species since hardly any marker in common to a closely related Triticum species could be identified.

The interval mapping method is widely used for the genetic mapping of quantitative trait loci (QTLs), though true resolution of quantitative variation into QTLs is hampered with this method. Separation of QTLs is troublesome, because single-QTL is models are fitted. Further, genotype-by-environment interaction, which is of great importance in many quantitative traits, can only be approached by separately analyzing the data collected in multiple environments. Here, we demonstrate for the first time a novel analytic approach (MQM mapping) that accommodates both the mapping of multiple QTLs and genotype-by-environment interaction. MQM mapping is compared to interval mapping in the mapping of QTLs for flowering time in Arabidopsis thaliana under various photoperiod and vernalization conditions.

BACKGROUND: The current system for financing effective mental health can improve or reduce the quality of care provided. The system of financing includes reimbursing the care-provider, a risk-adjustment system for the health insurer and personal payments by the patient. Care-providers, patients and health insurers are worried that the current system is detrimental to quality care. AIM: To find out whether the financing system can be improved by the introduction of a system which weighs the costs according to the intensity of the care provided and which is based on staging and profiling. METHOD: We performed a critical evaluation of the current financing system and we explored how staging and profiling could help to keep complex care affordable. RESULTS: The current model for financing mental health care does not involve staging and profiling. Reimbursement is based on diagnoses and on the number of minutes of care provided, no differentiation being made between the levels of expertise required. As a result it can become financially unattractive to administer the best possible treatment to patients with the greatest needs. Staging and profiling have the potential to create the best possible relationship between the level of care needed by the patient and the treatment costs incurred, because the factors that led to the previous imbalance have been removed. CONCLUSION: Staging and profiling can be introduced gradually at roughly the same pace as the national system of risk-adjustment is being introduced for health insurers.

Summary A computerized procedure to construct integrated genetic maps is presented. The computer program (Join Map) can handle raw data from F2s, backcrosses and recombinant inbred lines, as well as listed pair-wise recombination frequencies. The procedure is useful for combining linkage data that have been collected in different experiments; the result is a mathematical alignment of the distinct genetic maps. Data from single experiments can be dealt with as well. In view of the fast growing amount of linkage information for molecular markers, which is often being generated by different research groups, integrated maps provide useful information on the map position of genes and DNA markers. The procedure performs a sequential build-up of the map and, at each step, a numerical search for the best fitting order of markers. Weighted least squares is used for the estimation of map distances.

Background: Higher plants possess a large multigene family encoding secreted class III peroxidase (Prx) proteins. Peroxidases appear to be associated with plant disease resistance based on observations of induction during disease challenge and the presence or absence of isozymes in resistant vs susceptible varieties. Despite these associations, there is no evidence that allelic variation of peroxidases directly determines levels of disease resistance. Methodology/Principal Findings: The current study introduces a new strategy called Prx-Profiling. We showed that with this strategy a large number of peroxidase genes can be mapped on the barley genome. In order to obtain an estimate of the total number of Prx clusters we followed a re-sampling procedure, which indicated that the barley genome contains about 40 peroxidase gene clusters. We examined the association between the Prxs mapped and the QTLs for resistance of barley to homologous and heterologous rusts, and to the barley powdery mildew fungus. We report that 61% of the QTLs for partial resistance to P. hordei, 61% of the QTLs for resistance to B. graminis and 47% of the QTLs for non-host resistance to other Puccinia species co-localize with Prx based markers. Conclusions/Significance: We conclude that Prx-Profiling was effective in finding the genetic location of Prx genes on the barley genome. The finding that QTLs for basal resistance to rusts and powdery mildew fungi tend to co-locate with Prx clusters provides a base for exploring the functional role of Prx-related genes in determining natural differences in levels of basal resistance. © 2010 González et al.

Alternaria solani (Ellis and Martin) Sorauer, the causal agent of early blight (EB) disease, infects aerial parts of tomato at both seedling and adult plant stages. Resistant cultivars would facilitate a sustainable EB management. EB resistance is a quantitatively expressed character, a fact that has hampered effective breeding. In order to identify and estimate the effect of genes conditioning resistance to EB, a quantitative trait loci (QTL) mapping study was performed in F2 and F3 populations derived from the cross between the susceptible Solanum lycopersicum (syn. Lycopersicon esculentum) cv. `Solentos¿ and the resistant Solanum arcanum (syn. Lycopersicon peruvianum) LA2157 and genotyped with AFLP, microsatellite and SNP markers. Two evaluation criteria of resistance were used: measurements of EB lesion growth on the F2 plants in glasshouse tests and visual ratings of EB severity on foliage of the F3 lines in a field test. A total of six QTL regions were mapped on chromosomes 1, 2, 5¿7, and 9 with LOD scores ranging from 3.4 to 17.5. Three EB QTL also confer resistance to stem lesions in the field, which has not been reported before. All QTL displayed significant additive gene action; in some cases a dominance effect was found. Additive × additive epistatic interactions were detected between one pair of QTL. For two QTL, the susceptible parent contributed resistance alleles to both EB and stem lesion resistance. Three of the QTL showed an effect in all tests despite methodological and environmental differences.

The Mendelian analysis of genetic variation, available as induced mutants or as natural variation, requires a number of steps that are described in this chapter. These include the determination of the number of genes involved in the observed trait's variation, the determination of dominance relationships between alleles of the same locus, and epistatic interactions with related genetic variants. A new variant should be compared with previously identified genetic variants, which is most efficiently done by allelism tests in case of recessive mutants. In addition the locus should be localized on the genetic and physical map by linkage analysis. The mapping of mutant loci in Arabidopsis is facilitated by the genomic resources available in Arabidopsis and often consists of two steps, a crude and a fine mapping, the latter enabling the approximate location of a variant on the sequence map of Arabidopsis

Sixteen sweet potato varieties were evaluated for fresh storage root yield in 20 trials during 2000¿2001 for three seasons in four locations in Uganda. Of the 16 varieties, 11 were developed by farmers and five by a central breeding programme. The behaviour of the varieties was quantified in terms of wide adaptation (genotypic mean across trials), specific adaptation (genotypic predictions for specific locations) and stability (Shukla stability variance). With respect to all three aspects of yield behaviour, farmer varieties performed on average better than the official varieties. The results illustrate the potential that farmer varieties can have in the improvement of sweet potato in Uganda and other regions where high diversity of sweet potato landraces exists

A new method, REcombination Counting and ORDering (RECORD) is presented for the ordering of loci on genetic linkage maps. The method minimizes the total number of recombination events. The search algorithm is a heuristic procedure, combining elements of branch-and-bound with local reshuffling. Since the criterion we propose does not require intensive calculations, the algorithm rapidly produces an optimal ordering as well as a series of near-optimal ones. The latter provides insight into the local certainty of ordering along the map. A simulation study was performed to compare the performance of RECORD and JoinMap. RECORD is much faster and less sensitive to missing observations and scoring errors, since the optimisation criterion is less dependent on the position of the erroneous markers. In particular, RECORD performs better in regions of the map with high marker density. The implications of high marker densities on linkage map construction are discussed.

Breeding of cut and pot rose cultivars for efficient production under low-energy conditions in greenhouses will be facilitated by understanding the inheritance of vigour. To get insight into the genetic variation of vigour-related traits, a diploid rose population was employed for an evaluation study in greenhouses in The Netherlands and Denmark. For all the traits investigated the population showed a continuous quantitative variation as well as a considerable transgression. For most of the traits, the genetic variation found among the tested entries was highly significant and tended to be large in comparison to the effects of genotype by environment interaction. The heritability based on means of the traits was high and ranged from 68 to 92%. Strong simple correlations (r = 0.65 to 0.95) were found among the traits shoot length, leaf area, leaf dry weight, stem dry weight, total dry weight and growth rate. The total dry weight and leaf area are suggested to be good parameters for early selection of rose genotypes with vigorous growth under suboptimal growth conditions.

Species from productive and unproductive habitats differ inherently in their relative growth rate (RGR) and a wide range of correlated quantitative traits. We investigated the genetic basis of this trait complex, and specifically assessed whether it is under the control of just one or a few genes that can act as master switches by simultaneously affecting a range of traits in the complex. To address this problem, we crossed two Hordeum spontaneum lines originating from two habitats that differ in productivity. The F3 offspring, in which parental alleles are present in different combinations due to recombination and segregation, was analysed for RGR and its underlying components (leaf area ratio, unit leaf rate, photosynthesis, respiration), as well as a number of other physiological and morphological parameters. For this intra-specific comparison, we found a complex of positively and negatively correlated traits, which was quite similar to what is generally observed across species. A quantitative trait loci (QTL) analysis showed three major and one minor QTL for RGR. Most other variables of the growth-trait complex showed fewer QTLs that were typically scattered over various locations on the genome. Thus, at least in H. spontaneum, we found no evidence for regulation of the trait complex by one or two master switches. [KEYWORDS: Specific leaf area; Unit leaf rate; Quantitative trait loci; Trait complex; Photosynthesis]

Combining ecophysiological modelling and genetic mapping has increasingly received attention from researchers who wish to predict complex plant or crop traits under diverse environmental conditions. The potential for using this combined approach to predict flowering time of individual genotypes in a recombinant inbred line (RIL) population of spring barley (Hordeum vulgare L.) was examined. An ecophysiological phenology model predicts preflowering duration as affected by temperature and photoperiod, based on the following four input traits: f(o) (the minimum number of days to flowering at the optimum temperature and photoperiod), theta1 and theta2 (the development stages for the start and the end of the photoperiod-sensitive phase, respectively), and delta (the photoperiod sensitivity). The model-input trait values were obtained from a photoperiod-controlled greenhouse experiment. Assuming additivity of QTL effects, a multiple QTL model was fitted for the model-input traits using composite interval mapping. Four to seven QTL were identified for each trait. Each trait had at least one QTL specific to that trait alone. Other QTL were shared by two or all traits. Values of the model-input traits predicted for the RILs from the QTL model were fed back into the ecophysiological model. This QTL-based ecophysiological model was subsequently used to predict preflowering duration (d) for eight field trial environments. The model accounted for 72% of the observed variation among 94 RILs and 94% of the variation among the two parents across the eight environments, when observations in different environments were pooled. However, due to the low percentage (34-41%) of phenotypic variation accounted for by the identified QTL for three model-input traits (theta1, theta2 and delta), the QTL-based model accounted for somewhat less variation among the RILs than the model using original phenotypic input trait values. Nevertheless, days to flowering as predicted from the QTL-based ecophysiological model were highly correlated with days to flowering as predicted from QTL-models per environment for days to flowering per se. The ecophysiological phenology model was thus capable of extrapolating (QTL) information from one environment to another.

Associations between markers and complex quantitative traits were investigated in a collection of 146 modern two-row spring barley cultivars, representing the current commercial germ plasm in Europe. Using 236 AFLP markers, associations between markers were found for markers as far apart as 10 cM. Subsequently, for the 146 cultivars the complex traits mean yield, adaptability (Finlay-Wilkinson slope), and stability (deviations from regression) were estimated from the analysis of variety trial data. Regression of those traits on individual marker data disclosed marker-trait associations for mean yield and yield stability. Support for identified associations was obtained from association profiles, i.e., from plots of P-values against chromosome positions. In addition, many of the associated markers were located in regions where earlier QTL were found for yield and yield components. To study the oligogenic genetic base of the traits in more detail, multiple linear regression of the traits on markers was carried out, using stepwise selection. By this procedure, 18–20 markers that accounted for 40–58% of the variation were selected. Our results indicate that association mapping approaches can be a viable alternative to classical QTL approaches based on crosses between inbred lines, especially for complex traits with costly measurements.

High-density genetic linkage maps can be used for purposes such as fine-scale targeted gene cloning and anchoring of physical maps. However, their construction is significantly complicated by even relatively small amounts of scoring errors. Currently available software is not able to solve the ordering ambiguities in marker clusters, which inhibits the application of high-density maps. A statistical method named SMOOTH was developed to remove genotyping errors from genetic linkage data during the mapping process. The program SMOOTH calculates the difference between the observed and predicted values of data points based on data points of neighbouring loci in a given marker order. Highly improbable data points are removed by the program in an iterative process with a mapping algorithm that recalculates the map after cleaning. SMOOTH has been tested with simulated data and experimental mapping data from potato. The simulations prove that this method is able to detect a high amount of scoring errors and demonstrates that the program enables mapping software to successfully construct a very accurate high-density map. In potato the application of the program resulted in a reliable placement of nearly 1,000 markers in one linkage group.

A high-density genetic map with a number of anchor markers has been created to be used as a tool to dissect genetic variation in rose. Linkage maps for the diploid 94/1 population consisting of 88 individuals were constructed using a total of 520 molecular markers including AFLP, SSR, PK, RGA, RFLP, SCAR and morphological markers. Seven linkage groups, putatively corresponding to the seven haploid rose chromosomes, were identified for each parent, spanning 487 cM and 490 cM, respectively. The average length of 70 cM may cover more than 90% of the rose genome. An integrated map was constructed by incorporating the homologous parental linkage groups, resulting in seven linkage groups with a total length of 545 cM. The present linkage map is currently the most advanced map in rose with regard to marker density, genome coverage and with robust markers, giving good perspectives for QTL mapping and marker-assisted breeding in rose. The SSR markers, together with RFLP markers, provide good anchor points for future map alignment studies in rose and related species. Codominantly scored AFLP markers were helpful in the integration of the parental maps.

The genetic basis of phenotypic plasticity of relative growth rate (RGR), its components and associated morphological traits was studied in relation to nutrient limitation. In all, 140 F3 lines from a cross, made between two Hordeum spontaneum (wild barley) accessions sampled in Israel, were subjected to growth analysis under two nutrient levels. Quantitative trait loci (QTLs) were detected for RGR and three of its components, leaf area ratio (LAR), specific leaf area and leaf mass fraction (LMF). Indications for close linkage (potential pleiotropy) were found, for example, for LAR and LMF. An interesting case is on chromosome 6, at which QTLs for RGR and seed mass were detected in the same region. These QTLs had opposite additive effects, supporting earlier results that plants growing from lighter seeds had a higher RGR. Only two QTLs were significant under both nutrient conditions, suggesting large QTL ´ environment interactions for most traits. For 21 out of 26 QTLs, however, the additive genetic effect was of identical sign in both nutrient environments, but reached the significance threshold in only one of them. Nevertheless, some QTLs detected in one of the two environments had virtually no effect in the other, and QTLs for plasticity were detected for RGR, LAR and LMF, as well as for some morphological traits. QTLs with opposite effects under high and low nutrients were not found. Thus, at the genetic level, there was no evidence for a trade-off between faster growth at high versus low nutrient levels [KEYWORDS: QTL; plasticity; nutrient limitation; relative growth rate; wild barley; QTL ´ environment interaction]

Wild relatives of crop plants offer an attractive gene pool for cultivar improvement. We evaluated genetic and phenotypic variation for a set of 72 Israeli accessions of wild barley from 21 populations. These populations were grouped further into four ecotypes. In addition, environmental variables describing the local conditions for the populations were used to infer the environmental divergence. Genetic, phenotypic and environmental distances were estimated from the data and UPGMA dendrograms constructed. The results showed that genetic variation was larger between populations than within them, whereas for phenotypic measurements variation was larger within populations than between them. No significant correlation was found between genetic and phenotypic similarities, or between phenotypic and environmental similarities, whereas a weak correlation between genetic and environmental similarities was detected. Twenty-three AFLP-markers were identified to be ecotype specific. Chromosomal location was known for five of these markers. Four of the five ecotype specific markers were correlated with both phenotypic traits and environmental variables.

Lycopersicon peruvianum LA2172 is completely resistant to Oidium neolycopersici, the causal agent of tomato powdery mildew. Despite the large genetic distance between the cultivated tomato and L. peruvianum, fertile F-1 hybrids of L. esculentum cv. Moneymaker x L. peruvianum LA2172 were produced, and a pseudo-F-2 population was generated by mating F-1 half-sibs. The disease tests on the pseudo-F-2 population and two BC1 families showed that the resistance in LA2172 is governed by one dominant gene, designated as Ol-4. In the pseudo-F-2 population, distorted segregation was observed, and multi-allelic, single-locus markers were used to display different marker-allele configurations per locus. Parameters for both distortion and linkage between genetic loci were determined by maximum likelihood estimation, and the necessity of using multi-allelic, single-locus markers was illustrated. Finally, a genetic linkage map of chromosome 6 around the Ol-4 locus was constructed by using the pseudo-F-2 population.

In Northern Europe, dandelion populations consist solely of triploid or higher polyploid apomicts. Without a regular sexual cycle or lateral gene transmission, a clonal structure is expected for Taraxacum apomicts, although this was not found by compatibility analysis. In this study, we investigate whether this observation could be suported by performing independent tests based on data from hypervariable microsatellite markers as well as more conservative data based on allozymes and matrilinear cpDNA markers. In addition, population genetic methods were used to test departure from panmictic expectations, which is expected for clonal populations. Results indicated that many data sets, again, did not agree with expectations from clonal evolution because only small groups of genotypes exhibit no marker incompatibility. Population genetic analysis revealed that virtually all genotypes, but not individuals, agreed with random segregation and genotypic equilibria. Exceptions were genotypes with rare allozyme alleles or nearly identical microsatellite genotypes. Consequently, a population sample of apomictic dandelions essentially harbours genotypes that resulted from segregation and/or recombination and only a few genotypes that may have differentiated by somatic mutations [KEYWORDS: Taraxacum, apomixis, AFLPs, character incompatibility]

Genes contributing to the quantitative variation of a complex crop trait can be numerous. However, using existing approaches, the number of quantitative trait loci (QTL) detected for a trait is limited. Therefore, rather than looking for QTL for a complex trait itself, determining QTL for underlying component traits might give more information. In this study the potential of component analysis in QTL mapping of complex traits was examined using grain yield in spring barley as an example. Grain yield was divided into three components: number of spikes/m2, number of kernels/spike, and 1000-kernel weight. These traits were measured for individuals of a recombinant inbred-line population in field trials conducted over 2 years. By the use of an approximate multiple QTL model, one to eight QTL were detected for each trait in a year. Some QTL were mapped to similar positions in both years. Almost all QTL for yield were found at the position of or in close proximity to QTL for its component traits. A number of QTL for component traits were not detected when yield itself was subjected to QTL analysis. However, relative to the QTL for yield itself, all component-trait QTL did not explain the variation in yield better. The results in relation to the potential of using component analysis in studying complex crop traits are discussed.

The origin of plant breeding traces back to the dawn of agriculture and the domestication of plants, when nomadic man first became a settler. This started in those regions of the world where plants suitable for domestication, such as large grain grasses, grew in the wild. We envisage that the early

Advances in the use of molecular markers to elucidate the inheritance of quantitative traits enable the integration of genetic information on physiological traits into crop growth models. The objective of this study was to assess the ability of a crop growth model with QTL-based estimates of physiological input parameters to predict the yield of recombinant inbred lines (RILs) of barley. The model used predicts yield as spike biomass accumulated over the post-flowering period. We describe a two-stage procedure for predicting trait values from estimated additive and epistatic effects of QTLs. Values of physiological traits estimated by that procedure or measured in the field were used as input to the crop growth model. The output values (yield and shoot biomass) from the growth model using these two types of input values were highly correlated, indicating that QTL information can successfully replace measured input parameters. With the current crop growth model, however, both types of input values often resulted in large discrepancies between observed and predicted values. Improvement of performance may be achieved by incorporating physiological processes not yet included in the model. The prospects of using QTL-based predictions of model-input traits to identify new, high yielding barley genotypes are discussed.

By using a high-density AFLP marker linkage map, six QTLs for partial resistance to barley leaf rust (Puccinia hordei) isolate 1.2.1. have been identified in the RIL offspring of a cross between the partially resistant cultivar ’Vada’ and the susceptible line L94. Three QTLs were effective at the seedling stage, and five QTLs were effective at the adult plant stage. To study possible isolate specificity of the resistance, seedlings and adult plants of the 103 RILs from the cross L94×’Vada’ were also inoculated with another leaf rust isolate, isolate 24. In addition to the two QTLs that were effective against isolate 1.2.1. at the seedling stage, an additional QTL for seedling resistance to isolate 24 was identified on the long arm of chromosome 7. Of the eight detected QTLs effective at the adult plant stage, three were effective in both isolates and five were effective in only one of the two isolates. Only one QTL had a substantial effect at both the seedling and the adult plant stages. The expression of the other QTLs was developmental-stage specific. The isolate specificity of the QTLs supports the hypothesis of Parlevliet and Zadoks (1977) that partial resistance may be based on a minor-gene-for-minor-gene interaction.

Parents were selected from a well-characterised Arabidopsis recombinant inbred line (RIL) population based on (1) their phenotype for flowering time or (2) marker and QTL information that had been assessed previously. The F2 offspring obtained from pairs of selected RILs was analysed for these traits, and the results obtained with these two methods of selection were compared. Selection based on marker and QTL information gave approximately the same result as selection based on phenotype. The relative high heritability of flowering time in Arabidopsis facilitated successful phenotypical selection. The difference in selection result that was anticipated to be in favour of the marker-assisted approach was therefore not observed.

Due to the complexity of tetrasomic inheritance, mapping studies in potato (Solanum tuberosum L.) are generally conducted at the diploid level. In the present study we tested the feasibility of Bulked Segregant Analysis (BSA) using a tetraploid offspring for the identification of AFLP markers linked to the R2 allele, which confers race-specific resistance to Phytophthora infestans. Eleven bulk-specific AFLP markers, detected in fingerprints of 205 AFLP primer combinations, could be mapped in a linkage group encompassing the R2 locus. The efficiency of BSA at the tetraploid level, determined by the frequency of single-dose restriction fragments (SDRF), was much higher than expected on the basis of overall genetic dissimilarity between the parental clones. The fortuitous detection of AFLPs with linkage to the R2 allele is explained on the basis of specific genetic dissimilarity between cultivated potato and the chromosomal segment introgressed from S. demissum carrying the resistant R2 allele. AFLP markers common to those with linkage to R2 were visually recognized by their electrophoretic mobility in the AFLP fingerprint in a parental clone of a reference mapping population. Using these common AFLP markers we anchored the linkage group comprising the R2 allele to potato chromosome 4.

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.

Thirty one barley lines were used to investigate the agreement between three relationship measures: genetic similarities based on 681 AFLP-markers, coefficients of co-ancestry based on pedigree data, and generalised distance based on 25 morpological characters (morphological distance). Bootstrap analysis was used to estimate the accuracy of the correlation estimates. AFLP-based genetic similarities showed a poor-to-moderate correlation with the coefficients of co-ancestry within the core set of 25 European two-row spring barleys. Morphological distance was not significantly correlated with either genetic similarity or the coefficient of co-ancestry. The precision of all correlation-coefficient estimates, however, was low. The inclusion of two European winter barleys, two North American two-row spring barleys, and two North American six-row spring barleys in the AFLP-analysis resulted in a much stronger correlation between genetic similarity and the coefficient of co-ancestry. This suggests good opportunities for the use of AFLP-markers to assess genetic diversity by distinguishing between the major ecotypes of barley. Additionally, each of the eight primer combinations used in the AFLP-analysis was able to identify all 31 lines uniquely, showing the usefulness of AFLPs for cultivar identification. Because of the inaccuracy of the investigated relationship measures, resulting in low values of the correlation-coefficient estimates, prediction of the breeding behaviour of parent combinations may be improved by the use of a combination of relationship measures, thus decreasing the effect of their individual independent errors.

Barley (Hordeum vulgare L.) is one of the most extensively studied food crops in recent molecular research. More than 1000 molecular markers have been located on the barley genome by using five independent populations. For the present study, four segregation data sets, 'Proctor' × 'Nudinka', 'Igri' × 'Franka', 'Steptoe' × 'Morex', and 'Harrington' × TR306, were downloaded from the publicly available GrainGenes databank. Since 22% of the markers are common to at least two of the independent data sets, we were able to establish an integrated map using the computer package JOINMAP v2.0. The integrated map contains 898 markers, covers 1060 cM, and removes many large gaps present in the individual maps. Comparison of the integrated map with the individual maps revealed that the overall linear order of markers is in good agreement and that the integrated map is consistent with the component maps. No significant reordering of markers was found. This conservative property of the barley genome makes the integrated map reliable and successful. Except for chromosome 7 (5H), marker clustering was observed in the centromeric regions, probably owing to the centromeric suppression of recombination. Based on this integrated map, geneticists and breeders can choose their favourite markers in any region of interest of the barley genome. Key words : Hordeum vulgare, RFLP, integrated map.