Marker assisted selection with optimised contributions of the candidates to selection

The benefits of marker assisted selection (MAS) are evaluated under realistic assumptions in schemes where the genetic contributions of the candidates to selection are optimised for maximising the rate of genetic progress while restricting the accumulation of inbreeding. MAS schemes were compared with schemes where selection is directly on the QTL (GAS or gene assisted selection) and with schemes where genotype information is not considered (PHE or phenotypic selection). A methodology for including prior information on the QTL effect in the genetic evaluation is presented and the benefits from MAS were investigated when prior information was used. The optimisation of the genetic contributions has a great impact on genetic response but the use of markers leads to only moderate extra short-term gains. Optimised PHE did as well as standard truncation GAS (i.e. with fixed contributions) in the short-term and better in the long-term. The maximum accumulated benefit from MAS over PHE was, at the most, half of the maximum benefit achieved from GAS, even with very low recombination rates between the markers and the QTL. However, the use of prior information about the QTL effects can substantially increase genetic gain, and, when the accuracy of the priors is high enough, the responses from MAS are practically as high as those obtained with direct selection on the QTL.     

Marker assisted selection for the improvement of two antagonistic traits under mixed inheritance

A Monte Carlo simulation was used to investigate the potential of Marker Assisted Selection (MAS) in a multiple-trait situation. Only additive effects were considered. The base population was assumed to be in linkage equilibrium and, next, the population was managed over 15 discrete generations, 10 males and 50 females were chosen out of the 100 candidates of each sex. Performance for two traits was simulated with an overall heritability of a given trait equal to 0.25 or 0.10 and the overall genetic correlation between traits was generally equal to -0.4 except in one case where it was equal to 0. The model involved one biallelic QTL, accounting for 10 or 20% of the genetic variance of a given trait, plus polygenes. Initial allelic frequencies at the QTL were generally equal to 0.5 but in one case were equal to 0.1 and 0.9. A marker with 120 different alleles in the 60 founder parents was simulated in the vicinity of the QTL. Two values of the recombination rate between these two loci were considered, 0.10 and 0.02. The genetic evaluation was based on a multiple-trait BLUP animal model, accounting (MAS) or not (conventional BLUP) for marker information. Two sets of simulations were run: (1) a "missing data"case, with males having no record for one of the traits, and (2) a "secondary trait"case, with one trait having a weight in the aggregate genotype 4 times less than the other trait and the QTL acting only on this secondary trait. In the first set, evaluation methods were found to mainly affect the accuracy of overall genetic values prediction for the trait with missing data. In comparison with BLUP, MAS led to an extra overall genetic response for the trait with missing data, which was strongly penalised under the conventional BLUP, and to a deficit in response for the other trait. This more balanced evolution of the two traits was obtained, however, at the expense of the long-term overall cumulated response for the aggregate genotype, which was 1 to 2.5% lower than the one obtained under the conventional BLUP. In the second set of simulation, in the case of low initial frequency (0.1) of the QTL allele favourable to the secondary trait, MAS was found to be substantially more efficient to avoid losing this allele than BLUP only when the QTL had a large effect and the marker was close. More benefits should be expected from MAS with more specific applications, such as early selection of animals, or by applying dynamic procedures i.e. letting the respective weights to QTL and polygenic values in the selection criterion vary across generation.     

Marker assisted selection of bacterial blight resistance genes in rice

Bacterial leaf blight caused by Xanthomonas oryzae pv. oryzae is one of the most important diseases affecting rice production in Asia. We were interested in surveying rice genotypes that are popularly used in the Indian breeding program for conferring resistance to bacterial blight, using 11 STMS and 6 STS markers. The basis of selection of these DNA markers was their close linkage to xa5, xa13, and Xa21 genes and their positions on the rice genetic map relative to bacterial blight resistance genes. Eight lines were found to contain the xa5 gene while two lines contained Xa21 gene and none of the lines contained the xa13 gene with the exception of its near-isogenic line. Using the polymorphic markers obtained in the initial survey, marker-assisted selection was performed in the F3 population of a cross between IR-64 and IET-14444 to detect lines containing multiple resistance genes. Of the 59 progeny lines analyzed, eight lines contained both the resistance genes, xa5 and Xa4.     

The effect of using approximate gametic variance covariance matrices on marker assisted selection by BLUP

Under additive inheritance, the Henderson mixed model equations (HMME) provide an efficient approach to obtaining genetic evaluations by marker assisted best linear unbiased prediction (MABLUP) given pedigree relationships, trait and marker data. For large pedigrees with many missing markers, however, it is not feasible to calculate the exact gametic variance covariance matrix required to construct HMME. The objective of this study was to investigate the consequences of using approximate gametic variance covariance matrices on response to selection by MABLUP. Two methods were used to generate approximate variance covariance matrices. The first method (Method A) completely discards the marker information for individuals with an unknown linkage phase between two flanking markers. The second method (Method B) makes use of the marker information at only the most polymorphic marker locus for individuals with an unknown linkage phase. Data sets were simulated with and without missing marker data for flanking markers with 2, 4, 6, 8 or 12 alleles. Several missing marker data patterns were considered. The genetic variability explained by marked quantitative trait loci (MQTL) was modeled with one or two MQTL of equal effect. Response to selection by MABLUP using Method A or Method B were compared with that obtained by MABLUP using the exact genetic variance covariance matrix, which was estimated using 15 000 samples from the conditional distribution of genotypic values given the observed marker data. For the simulated conditions, the superiority of MABLUP over BLUP based only on pedigree relationships and trait data varied between 0.1% and 13.5% for Method A, between 1.7% and 23.8% for Method B, and between 7.6% and 28.9% for the exact method. The relative performance of the methods under investigation was not affected by the number of MQTL in the model.     

Marker-assisted selection of restored male-fertile Brassica napus plants using a set of dominant RAPD markers

Bulked segregant analysis was used to identify RAPD markers in oilseed rape (Brassica napus L.) that were linked to a male fertility restorer gene for Ogura cytoplasmic male sterility. After screening for polymorphisms using 960 primers, 14 RAPD markers were mapped to a 25 cM region including the restorer locus, a mapping population of 242 F₂ individuals being employed. The map was used to select 11 markers that were investigated for polymorphisms between the restorer donor line and 46 recipient lines. A set of four RAPD markers, one in coupling phase with the restorer allele and three with the non-restorer allele, which were informative in all 46 combinations, were used in marker assisted selection of plants homozygous for the restorer allele. A total of 906 homozygous restored plants were found among the 4605 BC₁F₂ plants analysed. Phenotypic data of a subset of the classified plants was compared with the RAPD data and the expected number of recombinants was calculated from the map data. A close correspondence between the expected and observed numbers of plants with a deviating phenotype was found. Thus, use of a set of dominant RAPD markers provides a way obtaining reliable data for marker-assisted selection.     

From crop domestication to super-domestication

Research related to crop domestication has been transformed by technologies and discoveries in the genome sciences as well as information-related sciences that are providing new tools for bioinformatics and systems' biology. Rapid progress in archaeobotany and ethnobotany are also contributing new knowledge to understanding crop domestication. This sense of rapid progress is encapsulated in this Special Issue, which contains 18 papers by scientists in botanical, crop sciences and related disciplines on the topic of crop domestication. One paper focuses on current themes in the genetics of crop domestication across crops, whereas other papers have a crop or geographic focus. One feature of progress in the sciences related to crop domestication is the availability of well-characterized germplasm resources in the global network of genetic resources centres (genebanks). Germplasm in genebanks is providing research materials for understanding domestication as well as for plant breeding. In this review, we highlight current genetic themes related to crop domestication. Impressive progress in this field in recent years is transforming plant breeding into crop engineering to meet the human need for increased crop yield with the minimum environmental impact - we consider this to be 'super-domestication'. While the time scale of domestication of 10 000 years or less is a very short evolutionary time span, the details emerging of what has happened and what is happening provide a window to see where domestication might - and can - advance in the future.     

Ultra-simple DNA extraction method for marker-assisted selection using microsatellite markers in rice

We prevent an ultra-simple DNA extraction method for microsatellite analysis of rice. Each extraction requires only one microtube, one disposable pipette tip, TE buffer and few pieces (about 5 mm) of rice leaf tissue. This is sufficient for 200 PCR reactions. The extract can be kept in the freezer for long-term storage. Also, DNA can be extracted from 200–300 individuals in a few hours. These features enabled us to perform rapid largescale seedling genotyping required for marker-assisted selection. We have also examined the applicability of this method for other PCR-based markers: RAPDs, nuclear STS, chloroplast STS and chloroplast microsatellites.     

Polymorphic simple sequence repeat markers in chloroplast genomes of Solanaceous plants

PCR-based markers were developed from mononucleotide simple-sequence repeats in the chloroplast genome of Nicotiana tabacum and applied to the analysis of genetic diversity. These markers were found to detect high levels of polymorphism at three taxonomic levels in Solanaceous plants. Of 36 chloroplast loci examined, 26 show some degree of polymorphism among potato accessions. Among a set of 30 tetraploid potato cultivars it is apparent that a single chloroplast haplotype is prevalent, presumably a result of the widespread use as a female parent of the imported US cultivar Rough Purple Chili in the latter half of the 19th century. Nonetheless, there is considerable chloroplast diversity in the cultivated potato, and it is clear that a large proportion of this variability has arisen through the use of wild or primitive cultivated species of potato in introgression programmes. This variability should be used in future breeding programmes. An examination of single accessions from 24 potato species, as well as representatives from tobacco and other members of the Solanaceae, reveals high levels of inter-specific chloroplast DNA variation. These data, and the ease of use and potential for multiplexing of these markers, suggest that cpSSRs will be of great utility in population genetics, germplasm management, evolutionary and phylogenetic studies as well as in, the analysis of material from introgression and somatic-fusion experiments. Interestingly, the polymorphism arising from one of the more-polymorphic chloroplast loci examined, does not originate solely from the SSR, and is due to variation in the copy number of two tandemly arrayed sequence elements. Received: 15 December 1998 / Accepted: 9 February 1999     

Bacterial leaf blight resistance genes (Xa21, xa13 and xa5) pyramiding through molecular marker assisted selection into rice cultivars

Marker assisted selection was employed to pyramid three bacterial blight resistance genes Xa21, xa13 and xa5 into high yielding susceptible rice cultivars ADT43 and ADT47. With the assistance of PCR markers, homozygous and heterozygous genotypes were identified in F₂ generation of two crosses (ADT43 × IRBB60 and ADT47 × IRBB60) and goodness of fit was tested. Eighty nine plants from F₃ generation of ADT43 × IRBB60 were also screened for resistance genes. The genotypes carrying resistance genes in different combinations were identified. The pyramided lines showed a wider spectrum and higher level of resistance against two Xoo isolates under field conditions.     

Seed paternity in flowering plants: an evolutionary perspective

The ultimate importance of paternal contributions to fitness and of post-pollination selection in flowering plants have remained elusive, largely because of the technical difficulty of assigning paternity. I review empirical studies that use heritable markers to investigate per-fruit seed paternity in natural populations and after experimental multiple-donor pollination. Thirty-one studies covering 23 species from 16 plant families document that in natural populations seeds from a single fruit are often fathered by multiple pollen donors (5 species from 5 families), that donors can differ significantly in seed-siring success (8 species from 6 families), that variation in pollen tube growth rates can be heritable (n = 1 out of 4 studies), that donor and recipient genotypes can simultaneously affect paternity (n = 2), and that temporal order of pollen deposition (n = 1) and environmental effects(n = 2) affect the outcome of pollen competition. These studies also investigate the role of post-pollination selection in the avoidance of inbreeding and for species boundaries. Most studies of male reproductive success in plants to date base on isozyme electrophoresis. The availability in the last decade of highly polymorphic molecular markers such as microsatellite DNA has been expected to open new possibilities to investigate competition and selection during the gametophytic phase. Yet, to date, there is still need for greater data wealth on seed paternity to test theories of sex allocation and to gain deeper understanding of floral trait evolution and of the evolutionary consequences of post-pollination selection in flowering plants.     

Changes in Variance Components of Flanking Marker Genotypes Under Varying Selection Intensities

Selection is practically ubiquitous during marker-QTL linkage analysis with an experimental population. Thus, it is necessary to investigate the impacts of selection upon linkage analyses in order to obtain unbiased estimates of QTL position and effect. In this article, by exploiting flanking markers through the widely applied half-sib design, we have developed the structures of three variance components, i.e., variance component between marker genotypes, polygenic variance component and recombinant variance component within marker genotypes. Changes in these variance components under varying selection intensities were investigated in this study to formulate the effects of selection on various variance components. Results showed clearly that all variance components presented were quite sensitive to changes in selection intensity. As selection intensity increased, all variance components declined by differing extents in a quadratic fashion. Comparatively speaking, the variance between marker genotypes decreased most drastically, followed by the polygenic variance within marker genotypes and then the recombinant variance within marker genotypes, which suggested a decrease of power for QTL linkage analysis. Therefore, steps should be taken to avoid as much as possible the presence of selection in real populations, so as to further eliminate the negative effects of selection on QTL linkage analysis.     

Genetic Analysis and Fine Mapping of Two Genes for Grain Shape and Weight in Rice

To identify genetic loci controlling grain weight, an elite indica rice variety, Baodali, with large grains was identified and used in this study. Its derived F2, F3 and BC2 F2 with another japonica rice variety Zhonghua 11 were used as mapping populations. Linkage analyses demonstrated that two genes controlling grain weight, designated as GW3 and GW6, were mapped to chromosome 3 and chromosome 6, respectively. Fine mapping delimited GW3 to a 122 kb physical distance between two sequence tagged site markers （WGWt6 and WGW19） containing 16 open reading frames annotated by The Institute for Genomic Research （http：//www.tigr.org）. GW6 was further mapped between two simple sequence repeat markers （RM7179 and RM3187）. These results are useful for both marker assisted selection of grain weight, and for further cloning of GW genes, which will contribute to the dissection of the molecular mechanism underlying grain weight in rice.     

侧翼标记基因型方差组分随不同选择强度的变化

从实际角度看,选择在用于标记-QTL连锁分析的实验群体中是较普遍存在的。为获得QTL位置及其效应的无偏估计,有必要研究选择对连锁分析实验所造成的影响。该文在侧翼标记情况下利用广泛使用的半同胞设计提出了3种方差组分,即标记组间方差、组内多基因方差和组内重组方差及其构成。着重从理论上研究了各方差组分在不同选择强度下的变化规律,阐明了选择对各方差组分的效应。结果表明,各方差组分对选择强度的变化很敏感。随着选择强度的增加,所有方差组分都以二次曲线方式有不同程度的下降。比较而言,标记组间方差组分下降最快,标记组内多基因方差下降较慢,组内重组方差下降最慢。这意味着QTL连锁分析功效的降低。所以在实际群体中应采取必要措施尽量避免选择的发生,从而消除选择对QTL连锁分析的负面效应。