Cassava (Manihot esculenta Crantz) is a clonally propagated staple food crop in the tropics. Genomic selection (GS) has been implemented at three breeding institutions in Africa to reduce cycle times. Initial studies provided promising estimates of predictive abilities. Here, we expand on previous analyses by assessing the accuracy of seven prediction models for seven traits in three prediction scenarios: cross-validation within populations, cross-population prediction and cross-generation prediction. We also evaluated the impact of increasing the training population (TP) size by phenotyping progenies selected either at random or with a genetic algorithm. Cross-validation results were mostly consistent across programs, with nonadditive models predicting of 10% better on average. Cross-population accuracy was generally low (mean = 0.18) but prediction of cassava mosaic disease increased up to 57% in one Nigerian population when data from another related population were combined. Accuracy across generations was poorer than within-generation accuracy, as expected, but accuracy for dry matter content and mosaic disease severity should be sufficient for rapid-cycling GS. Selection of a prediction model made some difference across generations, but increasing TP size was more important. With a genetic algorithm, selection of one-third of progeny could achieve an accuracy equivalent to phenotyping all progeny. We are in the early stages of GS for this crop but the results are promising for some traits. General guidelines that are emerging are that TPs need to continue to grow but phenotyping can be done on a cleverly selected subset of individuals, reducing the overall phenotyping burden.
Cassava (Manihot esculenta Crantz) production is currently under threat from cassava brown streak disease (CBSD), a disease that is among the seven most serious obstacles to world’s food security. Three issues are of significance for CBSD. Firstly, the virus associated with CBSD, has co-evolved with cassava outside its center of origin for at least 90 years. Secondly, that for the last 74 years, CBSD was only limited to the low lands. Thirdly, that most research has largely focused on CBSD epidemiology and virus diversity. Accordingly, this paper focuses on CBSD genetics and/or breeding and hence, presents empirical data generated in the past 11 years of cassava breeding in Uganda. Specifically, this paper provides: 1) empirical data on CBSD resistance screening efforts to identify sources of resistance and/or tolerance; 2) an update on CBSD resistance population development comprising of full-sibs, half-sibs and S1 families and their respective field performances; and 3) insights into chromosomal regions and genes involved in CBSD resistance based on genome wide association analysis. It is expected that this information will provide a foundation for harmonizing on-going CBSD breeding efforts and consequently, inform the future breeding interventions aimed at combating CBSD.
Cassava (Manihot esculenta Crantz) is an important security crop that faces severe yield loses due to cassava brown streak disease (CBSD). Motivated by the slow progress of conventional breeding, genetic improvement of cassava is undergoing rapid change due to the implementation of quantitative trait loci mapping, Genome-wide association mapping (GWAS), and genomic selection (GS). In this study, two breeding panels were genotyped for SNP markers using genotyping by sequencing and phenotyped for foliar and CBSD root symptoms at five locations in Uganda. Our GWAS study found two regions associated to CBSD, one on chromosome 4 which co-localizes with a Manihot glaziovii introgression segment and one on chromosome 11, which contains a cluster of nucleotide-binding site-leucine-rich repeat (NBS-LRR) genes. We evaluated the potential of GS to improve CBSD resistance by assessing the accuracy of seven prediction models. Predictive accuracy values varied between CBSD foliar severity traits at 3 months after planting (MAP) (0.27–0.32), 6 MAP (0.40–0.42) and root severity (0.31–0.42). For all traits, Random Forest and reproducing kernel Hilbert spaces regression showed the highest predictive accuracies. Our results provide an insight into the genetics of CBSD resistance to guide CBSD marker-assisted breeding and highlight the potential of GS to improve cassava breeding.
Cassava production in the central, southern and eastern parts of Africa is under threat by cassava brown streak virus (CBSV). Yield losses of up to 100% occur in cases of severe infections of edible roots. Easy illegal movement of planting materials across African countries, and long-range movement of the virus vector (Bemisia tabaci) may facilitate spread of CBSV to West Africa. Thus, effort to pre-emptively breed for CBSD resistance in W. Africa is critical. Genomic selection (GS) has become the main approach for cassava breeding, as costs of genotyping per sample have declined. Using phenotypic and genotypic data (genotyping-by-sequencing), followed by imputation to whole genome sequence (WGS) for 922 clones from National Crops Resources Research Institute, Namulonge, Uganda as a training population (TP), we predicted CBSD symptoms for 35 genotyped W. African clones, evaluated in Uganda. The highest prediction accuracy (r = 0.44) was observed for cassava brown streak disease severity scored at three months (CBSD3s) in the W. African clones using WGS-imputed markers. Optimized TPs gave higher prediction accuracies for CBSD3s and CBSD6s than random TPs of the same size. Inclusion of CBSD QTL chromosome markers as kernels, increased prediction accuracies for CBSD3s and CBSD6s. Similarly, WGS imputation of markers increased prediction accuracies for CBSD3s and for cassava brown streak disease root severity (CBSDRs), but not for CBSD6s. Based on these results we recommend TP optimization, inclusion of CBSD QTL markers in genomic prediction models, and the use of high-density (WGS-imputed) markers for CBSD predictions across population.
Cassava (Manihot esculenta Crantz) is a major source of dietary carbohydrates for >700 million people globally. However, its long breeding cycle has slowed the rate of genetic gain for target traits. This study aimed to asses genetic variation, the level of inbreeding, and trait correlations in genomic selection breeding cycles. We used phenotypic and genotypic data from the National Crops Resources Research Institute (NaCRRI) foundation population (Cycle 0, C0) and the progeny (Cycle 1, C1) derived from crosses of 100 selected C0 clones as progenitors, both to evaluate and optimize genomic selection. The highest broad‐sense heritability (H2 = 0.95) and narrow‐sense heritability (h2 = 0.81) were recorded for cassava mosaic disease severity and the lowest for root weight per plot (H2 = 0.06 and h2 = 0.00). We observed the highest genetic correlation (rg= 0.80) between cassava brown streak disease root incidence measured at seedling and clonal stages of evaluation, suggesting the usefulness of seedling data in predicting clonal performance for cassava brown streak root necrosis. Similarly, high genetic correlations were observed between cassava brown streak disease severity (rg= 0.83) scored at 3 and 6 mo after planting (MAP) and cassava mosaic disease, scored at 3 and 6 MAP (rg= 0.95), indicating that data obtained on these two diseases at 6 MAP would suffice. Population differentiation between C0 and C1 was not well defined, implying that the 100 selected progenitors of C1 captured the diversity in the C0. Overall, genetic gain for most traits were observed from C0 to C1.
Cassava brown streak disease (CBSD) caused by Cassava brown streak virus (CBSV) and Ugandan cassava brown streak virus (UCBSV) is a threat to food security in sub-Saharan Africa, where the disease persistently reduces overall root quality and quantity resulting in up to 100% yield losses. Complexities in CBSD symptom expression and the damage caused on leaves, stems and roots throughout the 12 months of cassava growth require that appropriate ways of categorizing genotype response and optimal stages of evaluation be identified. This study aimed at: 1) determining plot based heritability of CBSD based on symptom expression and 2) categorizing genotype resistance to CBSD based on symptom expression. Herein, 41 genotypes were evaluated for two years at Namulonge with an additional evaluation conducted across three locations. Evaluations were done at three, six, nine and twelve months after planting. Genotype responses to CBSD varied significantly. High broad sense heritability estimates of up to 0.81 (incidence) and 0.71 (severity) were obtained.Average disease severity scores had higher broad sense heritability estimates (0.53 and 0.65) than maximum disease severity scores (0.33 and 0.61) for root and foliar severities respectively. These findings are important in choosing an appropriate evaluation method for CBSD. Genotypes displayed differing CBSD responses in type, locality and severity of symptoms. This suggested that genotypes had differences in mechanisms of resistance that can be exploited in CBSD resistance breeding.
Cassava brown streak disease (CBSD) caused by the rapidly evolving cassava brown streak viruses (CBSVs), causes immense yield losses to the cassava value chain in eastern and southern Africa. Western Africa, another region that heavily depends on cassava is under eminent threat from CBSD. Resistance breeding is the best practical solution. However, complexities associated with CBSD resistance screening i.e., variable root sampling units, limit systematic attainment of genetic progress. Accordingly, we compared efficiency of five CBSD root necrosis assessment methods to guide selection: cassava brown streak disease root incidence (CBSDRi), cassava brown streak disease root severity (CBSDRs), cassava brown streak disease root severity computed as harmonic mean (CBSD-Harmonic), proportion-based root necrosis index (CBSD-proportion), and standardized root necrosis index (CBSD-standardized). The indexes (CBSD-proportion and CBSD-standardized) correct for variable sample size. We analyzed CBSD root necrosis data of 256 clones evaluated across 12 environments. Higher and variable standard errors were associated with root severity score 1 (no CBSD root necrosis). Lowest and highest plot-based heritability were respectively registered for CBSD-standardized (0.22) and CBSD-proportion (0.71). CBSDRs was only positively correlated with CBSDRi (r = 0.92) and CBSD-Harmonic (r = 0.97). Using best linear unbiased predictions (BLUPs), we ranked the top 15 CBSD resistant clones; only one clone (UG130014) featured in all the five assessment methods; two clones (UG130006 and UG120156) featured in four (CBSD-Harmonic, CBSDRi, CBSDRs, and CBSD-standardized); and five clones (UG120180, UG120063, UG130002, UG130033, and UG120183) featured in three methods (CBSD-Harmonic, CBSDRi, and CBSDRs). Influence of sample size was also quantified by sub-setting and analyzing CBSDRs data to have plots with at least 40 or 30 roots. Data stabilization was evident in plots with 30 roots. The significant influence of root sample sizes on overall ranking of clones, justifies the use of CBSD root necrosis indexes in early selection stages i.e., seedling and/or clonal trials, that are often characterized by high variations in roots assessed per plot. It is expected that this information will provide a foundation for harmonizing and/or optimizing on-going and future CBSD resistance breeding efforts.
Assembly of a training population (TP) is an important component of effective genomic selection-based breeding programs. In this study, we examined the power of diverse germplasm assembled from two cassava (Manihot esculenta Crantz) breeding programs in Tanzania at different breeding stages to predict traits and discover quantitative trait loci (QTL). This is the first genomic selection and genome-wide association study (GWAS) on Tanzanian cassava data. We detected QTL associated with cassava mosaic disease (CMD) resistance on chromosomes 12 and 16; QTL conferring resistance to cassava brown streak disease (CBSD) on chromosomes 9 and 11; and QTL on chromosomes 2, 3, 8, and 10 associated with resistance to CBSD for root necrosis. We detected a QTL on chromosome 4 and two QTL on chromosome 12 conferring dual resistance to CMD and CBSD. The use of clones in the same stage to construct TPs provided higher trait prediction accuracy than TPs with a mixture of clones from multiple breeding stages. Moreover, clones in the early Abbreviations: AYTKIB, advanced yield trial at Kibaha; BLUPs, best linear unbiased predictors; CBSD, cassava brown streak disease; CBSDRS, cassava brown streak disease root necrosis severity;
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