Aphis pomi (Hemiptera: Aphididae) population development, shoot characteristics, and antibiosis resistance in different apple genotypes

In high-value crops such as apple, Malus X domestica (Borkh.), insecticidal pest control is of high relevance. The use of resistant apple cultivars can increase the sustainability of pest management in apple orchards. Besides variation in plant chemistry that may influence plant resistance by antibiosis or antixenosis, plant growth characteristics also can affect plant susceptibility to pests such as aphids. Variable susceptibility to the apple aphid, Aphis pomi De Geer (Hemiptera: Aphididae), has been described for different apple cultivars. These observations were based on phenotypic surveys and no information on genetically based apple resistance to A. pomi is yet available. The objective of this study was to relate shoot growth characteristics with aphid population development, and to assess the genetic background of apple antibiosis-based resistance to A. pomi by quantitative trait loci (QTL) analysis. Aphid population development was repeatedly studied in the field in sleeve cages attached to 200 apple trees of different genotypes. Aphid population development was positively correlated to shoot length and growth, and it also was affected by climatic conditions. Indications for antibiosis-based resistance to A. pomi remained weak in the studied apple genotypes, and the only detected putative QTL on linkage group 11 of'Fiesta' apples was not stable for the different replications of the experiment. This lack of quantifiable resistance may be partly explained by environmental conditions related to aphid development in sleeve cages.     

Prospects for increasing yield in macadamia using component traits and genomics

Selection of candidate cultivars in macadamia requires extensive phenotypic measurements over many years and trials. In particular, yield traits such as nut-in-shell yield and kernel yield are economically vital characteristics and therefore guide the selection process for new cultivars. However, these traits can only be measured in mature trees, resulting in long generation intervals and slow rates of genetic gain. In addition, these traits are expensive to measure. Strategies to reduce the generation interval and increase the intensity of selection include using yield component traits, identification of markers associated with component traits, and genomic selection for yield. Yield component traits that contribute to resource availability for fruit formation include floral and nut characteristics. In this review, these traits will be investigated to estimate their relative importance in macadamia breeding and their heritability and correlations with yield. Furthermore, the usefulness of genome-wide association studies regarding yield component traits will be reviewed. Genetic-based breeding techniques could exploit this information to increase yield gains per breeding cycle and estimate the quantitative nature of yield traits. Genomic selection uses genome-wide molecular markers to predict the phenotype of individuals at an early age before maturity, thereby reducing the cycle time and increasing gain per unit time in plant breeding programmes. This review evaluates the potential for measurement of yield component traits, genome-wide association studies, and genomic selection to be employed in the Australian macadamia breeding programme to accelerate gains for nut yield.     

Improving breeding efficiency in potato using molecular and quantitative genetics

Key message

Potatoes are highly heterozygous and the conventional breeding of superior germplasm is challenging, but use of a combination of MAS and EBVs can accelerate genetic gain.

Abstract

Cultivated potatoes are highly heterozygous due to their outbreeding nature, and suffer acute inbreeding depression. Modern potato cultivars also exhibit tetrasomic inheritance. Due to this genetic heterogeneity, the large number of target traits and the specific requirements of commercial cultivars, potato breeding is challenging. A conventional breeding strategy applies phenotypic recurrent selection over a number of generations, a process which can take over 10 years. Recently, major advances in genetics and molecular biology have provided breeders with molecular tools to accelerate gains for some traits. Marker-assisted selection (MAS) can be effectively used for the identification of major genes and quantitative trait loci that exhibit large effects. There are also a number of complex traits of interest, such as yield, that are influenced by a large number of genes of individual small effect where MAS will be difficult to deploy. Progeny testing and the use of pedigree in the analysis can provide effective identification of the superior genetic factors that underpin these complex traits. Recently, it has been shown that estimated breeding values (EBVs) can be developed for complex potato traits. Using a combination of MAS and EBVs for simple and complex traits can lead to a significant reduction in the length of the breeding cycle for the identification of superior germplasm.     

The construction of a genetic linkage map of red raspberry (Rubus idaeus subsp. idaeus) based on AFLPs, genomic-SSR and EST-SSR markers

Breeding in raspberry is time-consuming due to the highly heterozygous nature of this perennial fruit crop, coupled with relatively long periods of juvenility. The speed and precision of raspberry breeding can be improved by genetic linkage maps, thus facilitating the development of diagnostic markers for polygenic traits and the identification of genes controlling complex phenotypes. A genetic linkage map (789 cM) of the red raspberry Rubus idaeus has been constructed from a cross between two phenotypically different cultivars; the recent European cultivar Glen Moy and the older North American cultivar Latham. SSR markers were developed from both genomic and cDNA libraries from Glen Moy. These SSRs, together with AFLP markers, were utilised to create a linkage map. In order to test the utility of the genetic linkage map for QTL analysis, morphological data based on easily scoreable phenotypic traits were collected. The segregation of cane spininess, and the root sucker traits of density and spread from the mother plant, was quantified in two different environments. These traits were analysed for significant linkages to mapped markers using MapQTL and were found to be located on linkage group 2 for spines and group 8 for density and diameter. The availability of co-dominant markers allowed heterozygosities to be calculated for both cultivars.     

Mapping quantitative physiological traits in apple (Malus × domestica Borkh.)

Efficient breeding and selection of high-quality apple cultivars requires knowledge and understanding of the underlying genetics. The availability of genetic linkage maps constructed with molecular markers enables the detection and analysis of major genes and quantitative trait loci contributing to the quality traits of a genotype. A segregating population of the cross between the apple varieties `Fiesta' (syn. `Red Pippin') and `Discovery' has been observed over three years at three different sites in Switzerland and data on growth habit, blooming behaviour, juvenile period and fruit quality has been recorded. QTL analyses were performed, based on a genetic linkage map consisting of 804 molecular markers and covering all 17 apple chromosomes. With the maximum likelihood based interval mapping method, the investigated complex traits could be dissected into a number of QTLs affecting the observed characters. Genomic regions participating in the genetic control of stem diameter, plant height increment, leaf size, blooming time, blooming intensity, juvenile phase length, time of fruit maturity, number of fruit, fruit size and weight, fruit flesh firmness, sugar content and fruit acidity were identified and compared with previously mapped QTLs in apple. Although `Discovery' fruit displayed a higher acid content, both acidity QTLs were attributed to the sweeter parent `Fiesta'. This indicated homozygosity at the acidity loci in `Discovery' preventing their detection in the progeny due to the lack of segregation.     

QTL mapping and breeding value estimation through pedigree-based analysis of fruit size and weight in four diverse peach breeding programs

The narrow genetic base of peach (Prunus persica L. Batsch) challenges efforts to accurately dissect the genetic architecture of complex traits. Standardized phenotypic assessment of pedigree-linked breeding germplasm and new molecular strategies and analytical approaches developed and conducted during the RosBREED project for enabling marker-assisted breeding (MAB) in Rosaceae crops has overcome several aspects of this challenge. The genetic underpinnings of fruit size (fruit equatorial diameter (FD)) and weight (fresh weight (FW)), two most important components of yield, were investigated using the pedigree-based analysis (PBA) approach under a Bayesian framework which has emerged as an alternative strategy to study the genetics of quantitative traits within diverse breeding germplasm across breeding programs. In this study, a complex pedigree with the common founder “Orange Cling” was identified and FD and FW data from 2011 and 2012 analyzed. A genetic model including genetic additive and dominance effects was considered, and its robustness was evaluated by using various prior and initial values in the Markov chain Monte Carlo procedure. Five QTLs were identified which accounted for up to 29 and 17 % of the phenotypic variation for FD and FW, respectively. Additionally, genomic breeding values were obtained for both traits, with accuracies >85 %. This approach serves as a model study for performing PBA across diverse pedigrees. By incorporating multiple breeding programs, the method and results presented support and highlight the ability of this strategy to identify genomic resources as targets for DNA marker development and subsequent MAB within each program.     

Integration of BpMADS4 on various linkage groups improves the utilization of the rapid cycle breeding system in apple

Rapid cycle breeding in apple is a new approach for the rapid introgression of agronomically relevant traits (e.g. disease resistances) from wild apple species into domestic apple cultivars (Malus × domestica Borkh.). This technique drastically shortens the long‐lasting juvenile phase of apple. The utilization of early‐flowering apple lines overexpressing the BpMADS4 gene of the European silver birch (Betula pendula Roth.) in hybridization resulted in one breeding cycle per year. Aiming for the selection of non‐transgenic null segregants at the end of the breeding process, the flower‐inducing transgene and the gene of interest (e.g. resistance gene) that will be introgressed by hybridization need to be located on different chromosomes. To improve the flexibility of the existing approach in apple, this study was focused on the development and characterization of eleven additional BpMADS4 overexpressing lines of four different apple cultivars. In nine lines, the flowering gene was mapped to different linkage groups. The differences in introgressed T‐DNA sequences and plant genome deletions post‐transformation highlighted the unique molecular character of each line. However, transgenic lines demonstrated no significant differences in flower organ development and pollen functionality compared with non‐transgenic plants. Hybridization studies using pollen from the fire blight‐resistant wild species accession Malus fusca MAL0045 and the apple scab‐resistant cultivar ‘Regia’ indicated that BpMADS4 introgression had no significant effect on the breeding value of each transgenic line.     

全基因组关联分析实现水稻粒型自然变异的分子解析

全基因组关联分析(GWAS)近年来被广泛应用于解析生物自然变异的遗传基础。但限于其遗传定位精度, 在水稻(Oryza sativa)遗传学研究中, 该方法尚无法取代传统的图位克隆法在克隆复杂性状调控基因中的作用。近期, 中国科学家在应用GWAS等大数据来克隆控制水稻粒长和粒重等复杂性状的QTL方面取得了新突破。     

Genetic transformation of perennial tropical fruits

Summary Genetic transformation provides the means for modifying single horticultural traits in perennial plant cultivars without altering their phenotype. This capability is particularly valuable for perennial plants and tree species in which development of new cultivars is often hampered by their long generation time, high levels of heterozygosity, nucellar embryony, etc. Most of these conditions apply to many tropical and subtropical fruit crops. Targeting specific gene traits is predicated upon the ability to regenerate elite selections of what are generally trees from cell and tissue cultures. The integrity of the clone would thereby remain unchanged except for the altered trait. This review provides an overview of the genetic transformation of perennial tropical and subtropical fruit crops, i.e., citrus (Citrus spp.), banana and plantain (Musa groups AAA, AAB, ABB, etc.), mango (Mangifera indica L.), pineapple (Ananas comosus L.), avocado (Persea americana Mill.), passion fruit (Passiflora edulis L.), longan (Dimocarpus longan Lour.), and litchi (Litchi chinensis Sonn.).     

Gene Tagging with Random Amplified Polymorphic DNA (RAPD) Markers for Molecular Breeding in Plants

Markers are of interest to plant breeders as a source of genetic information on crops and for use in indirect selection of traits to which the markers are linked. In the classic breeding approach, the markers were invariably the visible morphological and other phenotypic characters, and the breeders expended considerable effort and time in refining the crosses as the tight linkage or association of the desired characters with the obvious phenotypic characters was never unequivocally established. Furthermore, indirect selection for a trait using such morphological markers was not practical due to (1) a paucity of suitable markers, (2) the undesirable pleiotropic effects of many morphological markers on plant phenotype, and (3) the inability to score multiple morphological mutant traits in a single segregating population. With the advancement in molecular biology, the use of molecular markers in plant breeding has become very commonplace and has given rise to “molecular breeding”. Molecular breeding involves primarily “gene tagging”, followed by “marker-assisted selection” of desired genes or genomes. Gene tagging refers to the identification of existing DNA or the introduction of new DNA that can function as a tag or label for the gene of interest. In order for the DNA sequences to be conserved as a tag, important prerequisites exist. This review also summarizes the achievements in gene tagging that have been made over the last 7 to 8 years.     

Marker-Based Estimation of Heritability in Immortal Populations

Heritability is a central parameter in quantitative genetics, from both an evolutionary and a breeding perspective. For plant traits heritability is traditionally estimated by comparing within- and between-genotype variability. This approach estimates broad-sense heritability and does not account for different genetic relatedness. With the availability of high-density markers there is growing interest in marker-based estimates of narrow-sense heritability, using mixed models in which genetic relatedness is estimated from genetic markers. Such estimates have received much attention in human genetics but are rarely reported for plant traits. A major obstacle is that current methodology and software assume a single phenotypic value per genotype, hence requiring genotypic means. An alternative that we propose here is to use mixed models at the individual plant or plot level. Using statistical arguments, simulations, and real data we investigate the feasibility of both approaches and how these affect genomic prediction with the best linear unbiased predictor and genome-wide association studies. Heritability estimates obtained from genotypic means had very large standard errors and were sometimes biologically unrealistic. Mixed models at the individual plant or plot level produced more realistic estimates, and for simulated traits standard errors were up to 13 times smaller. Genomic prediction was also improved by using these mixed models, with up to a 49% increase in accuracy. For genome-wide association studies on simulated traits, the use of individual plant data gave almost no increase in power. The new methodology is applicable to any complex trait where multiple replicates of individual genotypes can be scored. This includes important agronomic crops, as well as bacteria and fungi.     

Assessment of Genotype Ranking in Long-term Biomass Production of Salix Based on Juvenile Plant Traits: Breeding Implications

Willow (Salix spp.) is among the most promising energy crops to be grown on agricultural land and breeding research to increase biomass yield of this perennial crop is performed in Europe and North America. Biomass willows are grown in short rotation and harvests are performed every 3 to 5 years (i.e., at 3- to 5-year cutting cycles) for a period of up to 25 years. However, breeding programs to improve long-term biomass yield are often relying on the results of short-term screening studies performed on juvenile plants. A pre-requisite for successful breeding of perennial energy crops is thus the identification of relevant juvenile plant traits indicative of long-term plant performance under field conditions. In this study a number of juvenile plant traits, measured at various Salix genotypes grown in a short-term experiment were evaluated in terms of their capacity to predict the long-term performance in biomass production after the first and second cutting cycle. The objective was to develop a simple model linking juvenile plant traits such as shoot biomass, total leaf area and leaf nitrogen (N) concentration to the long-term biomass productivity of field-grown plants. A two-component regression model combining juvenile shoot biomass and leaf N concentration provided the highest prediction accuracy (coefficients of determination around 0.8). The model based on two easy-to-measure juvenile plant traits clearly has implications for willow breeding programs. The implications for breeding are discussed in the light of the possibilities and limitations associated with the chosen approach.     

TGB: the tobacco genetics and breeding database

The germplasm of the genus Nicotiana contains more than 5,000 accessions and plays an important role in modern biological research. Tobacco can be used as a model system to develop methodologies for plant transformation and for investigating gene function. In order to develop the study of Nicotiana, a large quantity of data on germplasm, sequences, molecular markers and genetically modified tobacco was required for in-depth and systematic collation and research. It became necessary to establish a special database for tobacco genetics and breeding. The tobacco genetics and breeding (TGB, http://yancao.sdau.edu.cn/tgb) database was developed with the aim of bringing together tobacco genetics and breeding. The database has three main features: (1) a materials database with information on 1,472 Nicotiana germplasm accessions, as well as updated genomic and expressed sequence tag (EST) data available from the public database; (2) a molecular markers database containing a total of 12,388 potential intron polymorphisms 10,551 EST-simple sequence repeat (EST-SSR) and 66,297 genomic-SSR markers; and (3) an applications database with genetic maps and some genetically modified studies in tobacco. The TGB database also makes Basic Local Alignment Search Tool and primer designing tools publicly available. As far as can be ascertained, the TGB database is the first tobacco genetics and breeding database to be created, and all this comprehensive information will aid basic research into Nicotiana and other related plants. It will serve as an excellent resource for the online tobacco research community.     

A microRNA allele that emerged prior to apple domestication may underlie fruit size evolution

The molecular genetic mechanisms underlying fruit size remain poorly understood in perennial crops, despite size being an important agronomic trait. Here we show that the expression level of a microRNA gene (miRNA172) influences fruit size in apple. A transposon insertional allele of miRNA172 showing reduced expression associates with large fruit in an apple breeding population, whereas over‐expression of miRNA172 in transgenic apple significantly reduces fruit size. The transposon insertional allele was found to be co‐located with a major fruit size quantitative trait locus, fixed in cultivated apples and their wild progenitor species with relatively large fruit. This finding supports the view that the selection for large size in apple fruit was initiated prior to apple domestication, likely by large mammals, before being subsequently strengthened by humans, and also helps to explain why signatures of genetic bottlenecks and selective sweeps are normally weaker in perennial crops than in annual crops.     

Quantitative trait loci controlling plant architectural traits in cotton

Cotton plant architecture is an important characteristic influencing the suitability of specific cotton varieties in cultivation, fiber yield and quality. However, complex multigenic relationships and substantial genotype–environment interaction underlie plant architecture, and will hinder the efficient improvement of these traits in conventional cotton breeding programs. An enhanced understanding of the molecular-genetic regulation of plant morphological developmental can aid in the modification of agronomically relevant traits. In this study, an interspecific Gossypium hirsutum and Gossypium barbadense BC₁ population was used to identify QTL associated with plant architectural traits. Twenty-six single QTL were identified for seven plant architecture traits. The phenotypic variation explained by an individual QTL ranged from 9.56% to 44.57%. In addition, 11 epistatic QTL for fruit branch angle (FBA), plant height (PH), main-stem leaf size (MLS), and fruiting branch internode length (FBI) explained 2.28–15.34% of the phenotypic variation in these traits. The majority of the interactions (60%) occurred between markers linked to QTL influencing the same traits. The QTL detected in this study are expected to be valuable in future breeding programs to develop cultivars exhibiting desirable cotton architecture.     

Applications of Molecular Markers in Fruit Crops for Breeding Programs—A Review

Selection and use of molecular markers for evaluation of DNA polymorphism in plants are couple of the most important approaches in the field of molecular genetics. The assessment of genetic diversity using morphological markers is not sufficient due to little differentiating traits among the species, genera or their individuals. Morphological markers are not only highly influenced by environmental factors but skilled assessment is also prerequisite to find the variations in plant genetic resources. Therefore, molecular markers are considered as efficient tools for detailed DNA based characterization of fruit crops. Molecular markers provide new directions to the efforts of plant breeders particularly in genetic variability, gene tags, gene localization, taxonomy, genetic diversity, phylogenetic analysis and also play an important role to decrease the time required for development of new and excellent cultivars. The success of molecular markers technology in genetic improvement programs depends on the close relationship among the plant breeders, biotechnologists, skilled manpower and good financial support. The present review describes application and success of molecular markers technology used for genetic improvement in different fruit crops.     

Inducing drought tolerance in plants: Recent advances

Undoubtedly, drought is one of the prime abiotic stresses in the world. Crop yield losses due to drought stress are considerable. Although a variety of approaches have been used to alleviate the problem of drought, plant breeding, either conventional breeding or genetic engineering, seems to be an efficient and economic means of tailoring crops to enable them to grow successfully in drought-prone environments. During the last century, although plant breeders have made ample progress through conventional breeding in developing drought tolerant lines/cultivars of some selected crops, the approach is, in fact, highly time-consuming and labor- and cost-intensive. Alternatively, marker-assisted breeding (MAB) is a more efficient approach, which identifies the usefulness of thousands of genomic regions of a crop under stress conditions, which was, in reality, previously not possible. Quantitative trait loci (QTL) for drought tolerance have been identified for a variety of traits in different crops. With the development of comprehensive molecular linkage maps, marker-assisted selection procedures have led to pyramiding desirable traits to achieve improvements in crop drought tolerance. However, the accuracy and preciseness in QTL identification are problematic. Furthermore, significant genetic × environment interaction, large number of genes encoding yield, and use of wrong mapping populations, have all harmed programs involved in mapping of QTL for high growth and yield under water limited conditions. Under such circumstances, a transgenic approach to the problem seems more convincing and practicable, and it is being pursued vigorously to improve qualitative and quantitative traits including tolerance to biotic and abiotic stresses in different crops. Rapid advance in knowledge on genomics and proteomics will certainly be beneficial to fine-tune the molecular breeding and transformation approaches so as to achieve a significant progress in crop improvement in future. Knowledge of gene regulation and signal transduction to generate drought tolerant crop cultivars/lines has been discussed in the present review. In addition, the advantages and disadvantages as well as future prospects of each breeding approach have also been discussed.     

Sources of resistance and susceptibility to Septoria tritici blotch of wheat

An association genetics analysis was conducted to investigate the genetics of resistance to Septoria tritici blotch, caused by the fungus Zymoseptoria tritici (alternatively Mycosphaerella graminicola), in cultivars and breeding lines of wheat (Triticum aestivum) used in the UK between 1860 and 2000. The population was tested with Diversity Array Technology (DArT) and simple‐sequence repeat (SSR or microsatellite) markers. The lines formed a single population with no evidence for subdivision, because there were several common ancestors of large parts of the pedigree. Quantitative trait loci (QTLs) controlling Septoria resistance were postulated on 11 chromosomes, but 38% of variation was not explained by the identified QTLs. Calculation of best linear unbiased predictions (BLUPs) identified lineages of spring and winter wheat carrying different alleles for resistance and susceptibility. Abundant variation in Septoria resistance may be exploited by crossing well‐adapted cultivars in different lineages to achieve transgressive segregation and thus breed for potentially durable quantitative resistance, whereas phenotypic selection for polygenic quantitative resistance should be effective in breeding cultivars with increased resistance. The most potent allele reducing susceptibility to Septoria, on chromosome arm 6AL, was associated with reduced leaf size. Genes which increase susceptibility to Septoria may have been introduced inadvertently into UK wheat breeding programmes from cultivars used to increase yield, rust resistance and eyespot resistance between the 1950s and 1980s. This indicates the need to consider trade‐offs in plant breeding when numerous traits are important and to be cautious about the use of non‐adapted germplasm.     

Understanding the classics: the unifying concepts of transgressive segregation,inbreeding depression and heterosis and their central relevance for crop breeding

Transgressive segregation and heterosis are the reasons that plant breeding works. Molecular explanations for both phenomena have been suggested and play a contributing role. However, it is often overlooked by molecular genetic researchers that transgressive segregation and heterosis are most simply explained by dispersion of favorable alleles. Therefore, advances in molecular biology will deliver the most impact on plant breeding when integrated with sources of heritable trait variation – and this will be best achieved within a quantitative genetics framework. An example of the power of quantitative approaches is the implementation of genomic selection, which has recently revolutionized animal breeding. Genomic selection is now being applied to both hybrid and inbred crops and is likely to be the major source of improvement in plant breeding practice over the next decade. Breeders’ ability to efficiently apply genomic selection methodologies is due to recent technology advances in genotyping and sequencing. Furthermore, targeted integration of additional molecular data (such as gene expression, gene copy number and methylation status) into genomic prediction models may increase their performance. In this review, we discuss and contextualize a suite of established quantitative genetics themes relating to hybrid vigour, transgressive segregation and their central relevance to plant breeding, with the aim of informing crop researchers outside of the quantitative genetics discipline of their relevance and importance to crop improvement. Better understanding between molecular and quantitative disciplines will increase the potential for further improvements in plant breeding methodologies and so help underpin future food security.