Cassava retting ability and textural attributes of fufu for demand-driven cassava breeding.

BACKGROUND: Cassava retting ability and the textural qualities of cooked fufu are important quality traits. Cassava retting is a complex process in which soaking causes tissue breakdown, starch release, and softening. The rate at which various traits linked to it evolve varies greatly during fufu processing. According to the literature, there is no standard approach for determining retting ability. The retting indices and textural properties of fufu were measured using both manual and instrumental approaches. RESULTS: Different protocols were developed to classify 64 and 11 cassava genotypes into various groups based on retting ability and textural qualities, respectively. The retting protocols revealed considerable genetic dissimilarities in genotype classification: foaming ability and water clarity should be measured at 24 h, while penetrometer, hardness, turbidity, pH, and total titratable acidity data are best collected after 36 h. The stepwise regression model revealed that pH, foaming ability, and dry matter content are the best multivariates (with the highest R2) for predicting cassava retting. These predictors were used to develop an index for assessing the retting ability of cassava genotypes. The retting index developed showed a significant relationship with dry matter content and fufu yield. The study also showed significant correlations between instrumental cohesiveness and sensory smoothness (r = -0.75), moldability (r = -0.62), and stretchability (r = 0.78). Instrumental cohesiveness can correctly estimate fufu smoothness (R2 = 0.56, P = 0.008) and stretchability (R2 = 0.60, P = 0.005). CONCLUSION: pH, foaming ability, and dry matter content are the best traits for predicting cassava retting ability, while instrumental cohesiveness can effectively estimate fufu smoothness and stretchability. © 2024 The Authors. Journal of The Science of Food and Agriculture published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.

Content and distribution of cyanogenic compounds in cassava roots and leaves in association with physiological age.

BACKGROUND: Cassava roots are widely consumed in tropical regions of Asia, Africa, and Latin America. Although the protein, vitamin, carotenoid, and mineral content in the leaves makes them a nutritionally attractive option, their consumption is limited due to their high levels of cyanogenic compounds (CCs). In this study, the CC content in different parts of the plant (leaves, storage root cortex, and parenchyma) was assessed at harvest for 50 landrace genotypes representative of cassava diversity in Latin America. The changes in CC in leaves at different physiological ages (3, 6, 9, and 11 months after planting) were also investigated. RESULTS: The average CC was higher in the cortex (804 ppm) and leaves (655 ppm) than in root parenchyma (305 ppm). Genotypes from different regions of Latin America, as identified by seven genetic diversity groups, differed significantly in CC levels. The Andean and Amazon groups had, respectively, the lowest (P = 0.0008) and highest (P < 0.0001) CC levels in all three parts of the plants. Cyanogenic compound concentrations were higher in leaves from young plants (P < 0.0001) and decreased with increasing physiological age. CONCLUSION: The results help to guide the selection of parental lines with low CC levels for breeding and to contribute to the expanded use of cassava and its by-products for food and feed. Cassava for fresh consumption, especially, requires varieties with low total CC content, especially in the root cortex and parenchyma. COL1108 (204, 213, and 174 ppm, respectively, in the parenchyma, cortex, and leaves) and PER297 (83, 238, and 299 ppm, respectively, in the parenchyma, cortex, and leaves) can fulfill this requirement. © 2023 The Authors. Journal of The Science of Food and Agriculture published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.

Varietal impact on women's labour, workload and related drudgery in processing root, tuber and banana crops: focus on cassava in sub-Saharan Africa.

Roots, tubers and cooking bananas are bulky and highly perishable. In Africa, except for yams, their consumption is mainly after transport, peeling and cooking in the form of boiled pieces or dough, a few days after harvest. To stabilize and better preserve the products and, in the case of cassava, release toxic cyanogenic glucosides, a range of intermediate products have been developed, mainly for cassava, related to fermentation and drying after numerous processing operations. This review highlights, for the first time, the impact of genotypes on labour requirements, productivity and the associated drudgery in processing operations primarily carried out by women processors. Peeling, soaking/grinding/fermentation, dewatering, sieving and toasting steps were evaluated on a wide range of new hybrids and traditional landraces. The review highlights case studies of gari production from cassava. The results show that, depending on the genotypes used, women's required labour can be more than doubled and even the sum of the weights transported along the process can be up to four times higher for the same quantity of end product. Productivity and loads carried between each processing operation are highly influenced by root shape, ease of peeling, dry matter content and/or fiber content. Productivity and the often related experienced drudgery are key factors to be considered for a better acceptance of new genotypes by actors in the value-addition chain, leading to enhanced adoption and ultimately to improved livelihoods for women processors. © 2023 The Authors. Journal of The Science of Food and Agriculture published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.

East African highland cooking banana: towards an efficient selection of hybrids with user-preferred food quality traits.

BACKGROUND: Determinants of culinary qualities of East African highland cooking bananas (EAHCB) are not well known. This constrains the inclusion of user-preferred traits in breeding. The present study aimed to quantify key indicators of user-preferred characteristics to enable selection of acceptable hybrids. RESULTS: Qualitative characteristics that drive preference were big bunches (15-34 kg), long straight/slightly curved fingers (12-23 cm), yellowness and soft texture. Descriptive sensory analysis of the intensity of colour and texture the 23 genotypes revealed that landraces Kibuzi, Mbwazirume, Nakitembe and Mpologoma had higher intensity of yellowness and lower intensity of hardness (softer) and a low score (≤ 1.0) of astringency taste. A preference test showed that they had higher acceptability scores. Biochemical, instrumental and sensory data revealed correlations between sensory firmness and instrumental hardness (r = 0.5), sensory firmness and amylopectin (r = -0.54), suggesting that qualitative descriptions can be predicted by instrumental and biochemical indicators. Significant (P < 0.05) variations in amylose and total starch content were observed in different varieties. Moderate correlations between instrumental hardness and firmness in mouth (r = 0.55), cohesiveness and firmness in the mouth (r = 0.57), and adhesiveness and firmness in the mouth (r = 0.64) were observed. Surprisingly, carotenoids content was not correlated with yellowness in cooked matooke. However, positive correlations were observed between chroma (b*) parameters of raw matooke and sensorial assessed color on cooked samples. CONCLUSION: Qualitative characteristis; the bunch, pulp colour and texture; that drive users-preference in the EAHCB were quantified, paving way for breeders to use them to select genotypes with these attributes early in the breeding process. © 2023 The Authors. Journal of The Science of Food and Agriculture published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.

Characterizing quality traits of boiled yam: texture and taste for enhanced breeding efficiency and impact.

BACKGROUND: Boiled yam key quality attributes typical for West African consumers are that it is crumbly, easy to break and has a sweet taste. New yam varieties are being developed but high- or medium-throughput tools to assess the required quality traits and their range of acceptance are limited. This study assessed the acceptance thresholds of these quality attributes and established predictive models for screening yam varieties that meet the required consumer preferences. RESULTS: Overall liking was associated with sweet taste, crumbliness and easiness to break (r-values 0.502, 0.291 and -0.087, respectively). These parameters and selected biophysical parameters highly discriminated the boiled yam varieties. Crumbly texture and easiness to break were well predicted by penetration force and dry matter, whereas sweet taste were well predicted by dry matter and sugar intensity. A high crumbliness and sweet taste are preferred (sensory scores above 6.19 and 6.22 for crumbly and sweet taste, respectively, on a 10 cm unstructured line scale), while a too high easiness to break is disliked (sensory scores ranging from 4.72 to 7.62). Desirable biophysical targets were between 5.1 and 7.1 N for penetration force, dry matter around 39% and sugar intensity below 3.62 g 100 g-1 . Some improved varieties fulfilled the acceptable thresholds, and screening was improved through deviation from the optimum. CONCLUSION: Acceptance thresholds and deviation from optimum for boiled yam assessed through the instrumental measurements are promising tools for yam breeders. © 2023 The Authors. Journal of The Science of Food and Agriculture published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.

Field-transcriptome analyses reveal developmental transitions during flowering in cassava (Manihot esculenta Crantz).

KEY MESSAGE: We characterized genes that function in the photoperiodic flowering pathway in cassava. Transcriptome analysis of field-grown plants revealed characteristic expression patterns of these genes, demonstrating that field-grown cassava experiences two distinct developmental transitions. Cassava is an important crop for both edible and industrial purposes. Cassava develops storage roots that accumulate starch, providing an important source of staple food in tropical regions. To facilitate cassava breeding, it is important to elucidate how flowering is controlled. Several important genes that control flowering time have been identified in model plants; however, comprehensive characterization of these genes in cassava is still lacking. In this study, we identified genes encoding central flowering time regulators and examined these sequences for the presence or absence of conserved motifs. We found that cassava shares conserved genes for the photoperiodic flowering pathway, including florigen, anti-florigen and its associated transcription factor (GIGANTEA, CONSTANS, FLOWERING LOCUS T, CENTRORADIALIS/TERMINAL FLOWER1 and FD) and florigen downstream genes (SUPRESSOR OF OVEREXPRESSION OF CONSTANS1 and APETALA1/FRUITFUL). We conducted RNA-seq analysis of field-grown cassava plants and characterized the expression of flowering control genes. Finally, from the transcriptome analysis we identified two distinct developmental transitions that occur in field-grown cassava.

Fifty years of a public cassava breeding program: evolution of breeding objectives, methods, and decision-making processes.

This paper reviews and analyzes key features from cassava breeding at the International Center for Tropical Agriculture (CIAT) over 50 years and draws lessons for public breeding efforts broadly. The breeding team, jointly with national program partners and the private processing sector, defined breeding objectives and guiding business plans. These have evolved through the decades and currently focus on four global product profiles. The recurrent selection method also evolved and included innovations such as estimation of phenotypic breeding values, increasing the number of locations in the first stage of agronomic evaluations, gradual reduction of the duration of breeding cycles (including rapid cycling for high-heritability traits), the development of protocols for the induction of flowering, and the introduction of genome-wide predictions. The impact of cassava breeding depends significantly on the type of target markets. When roots are used for large processing facilities for starch, animal feeding or ethanol production (such as in SE Asia), the adoption of improved varieties is nearly universal and productivity at the regional scale increases significantly. When markets and relevant infrastructure are weak or considerable proportion of the production goes for local artisanal processing and on-farm consumption, the impact has been lower. The potential of novel breeding tools needs to be properly assessed for the most effective allocation of resources. Finally, a brief summary of challenges and opportunities for the future of cassava breeding is presented. The paper describes multiple ways that public and private sector breeding programs can learn from each other to optimize success.

Identification and validation of mutation points associated with waxy phenotype in cassava.

BACKGROUND: The granule-bound starch synthase I (GBSSI) enzyme is responsible for the synthesis of amylose, and therefore, its absence results in individuals with a waxy starch phenotype in various amylaceous crops. The validation of mutation points previously associated with the waxy starch phenotype in cassava, as well as the identification of alternative mutant alleles in the GBSSI gene, can allow the development of molecular-assisted selection to introgress the waxy starch mutation into cassava breeding populations. RESULTS: A waxy cassava allele has been identified previously, associated with several SNPs. A particular SNP (intron 11) was used to develop SNAP markers for screening heterozygote types in cassava germplasm. Although the molecular segregation corresponds to the expected segregation at 3:1 ratio (dominant gene for the presence of amylose), the homozygotes containing the SNP associated with the waxy mutation did not show waxy phenotypes. To identify more markers, we sequenced the GBSS gene from 89 genotypes, including some that were segregated from a cross with a line carrying the known waxy allele. As a result, 17 mutations in the GBSSI gene were identified, in which only the deletion in exon 6 (MeWxEx6-del-C) was correlated with the waxy phenotype. The evaluation of mutation points by discriminant analysis of principal component analysis (DAPC) also did not completely discriminate the waxy individuals. Therefore, we developed Kompetitive Allele Specific PCR (KASP) markers that allowed discrimination between WX and wx alleles. The results demonstrated the non-existence of heterozygous individuals of the MeWxEx6-del-C deletion in the analyzed germplasm. Therefore, the deletion MeWxEx6-del-C should not be used for assisted selection in genetic backgrounds different from the original source of waxy starch. Also, the alternative SNPs identified in this study were not associated with the waxy phenotype when compared to a panel of accessions with high genetic diversity. CONCLUSION: Although the GBSSI gene can exhibit several mutations in cassava, only the deletion in exon 6 (MeWxEx6-del-C) was correlated with the waxy phenotype in the original AM206-5 source.

Cassava breeding and agronomy in Asia: 50 years of history and future directions.

In Asia, cassava (Manihot esculenta) is cultivated by more than 8 million farmers, driving the rural economy of many countries. The International Center for Tropical Agriculture (CIAT), in partnership with national agricultural research institutes (NARIs), instigated breeding and agronomic research in Asia, 1983. The breeding program has successfully released high-yielding cultivars resulting in an average yield increase from 13.0 t ha-1 in 1996 to 21.3 t ha-1 in 2016, with significant economic benefits. Following the success in increasing yields, cassava breeding has turned its focus to higher-value traits, such as waxy cassava, to reach new market niches. More recently, building resistance to invasive pests and diseases has become a top priority due to the emergent threat of cassava mosaic disease (CMD). The agronomic research involves driving profitability with advanced technologies focusing on better agronomic management practices thereby maintaining sustainable production systems. Remote sensing technologies are being tested for trait discovery and large-scale field evaluation of cassava. In summary, cassava breeding in Asia is driven by a combination of food and market demand with technological innovations to increase the productivity. Further, exploration in the potential of data-driven agriculture is needed to empower researchers and producers for sustainable advancement.

Conventional breeding, marker-assisted selection, genomic selection and inbreeding in clonally propagated crops: a case study for cassava.

Consolidates relevant molecular and phenotypic information on cassava to demonstrate relevance of heterosis, and alternatives to exploit it by integrating different tools. Ideas are useful to other asexually reproduced crops. Asexually propagated crops offer the advantage that all genetic effects can be exploited in farmers' production fields. However, non-additive effects complicate selection because, while influencing the performance of the materials under evaluation, they cannot be transmitted efficiently to the following cycle of selection. Cassava can be used as a model crop for asexually propagated crops because of its diploid nature and the absence of (known) incompatibility effects. New technologies such as genomic selection (GS), use of inbred progenitors based on doubled haploids and induction of flowering can be employed for accelerating genetic gains in cassava. Available information suggests that heterosis, non-additive genetic effects and within-family variation are relatively large for complex traits such as fresh root yield, moderate for dry matter or starch content in the roots, and low for defensive traits (pest and disease resistance) and plant architecture. The present article considers the potential impact of different technologies for maximizing gains for key traits in cassava, and highlights the advantages of integrating them. Exploiting heterosis would be optimized through the implementation of reciprocal recurrent selection. The advantages of using inbred progenitors would allow shifting the current cassava phenotypic recurrent selection method into line improvement, which in turn would allow designing outstanding hybrids rather than finding them by trial and error.

Provitamin A accumulation in cassava (Manihot esculenta) roots driven by a single nucleotide polymorphism in a phytoene synthase gene.

Cassava (Manihot esculenta) is an important staple crop, especially in the arid tropics. Because roots of commercial cassava cultivars contain a limited amount of provitamin A carotenoids, both conventional breeding and genetic modification are being applied to increase their production and accumulation to fight vitamin A deficiency disorders. We show here that an allelic polymorphism in one of the two expressed phytoene synthase (PSY) genes is capable of enhancing the flux of carbon through carotenogenesis, thus leading to the accumulation of colored provitamin A carotenoids in storage roots. A single nucleotide polymorphism present only in yellow-rooted cultivars cosegregates with colored roots in a breeding pedigree. The resulting amino acid exchange in a highly conserved region of PSY provides increased catalytic activity in vitro and is able to increase carotenoid production in recombinant yeast and Escherichia coli cells. Consequently, cassava plants overexpressing a PSY transgene produce yellow-fleshed, high-carotenoid roots. This newly characterized PSY allele provides means to improve cassava provitamin A content in cassava roots through both breeding and genetic modification.

Cassava root crown phenotyping using three-dimension (3D) multi-view stereo reconstruction.

Phenotypic analysis of cassava root crowns (CRCs) so far has been limited to visual inspection and very few measurements due to its laborious process in the field. Here, we developed a platform for acquiring 3D CRC models using close-range photogrammetry for phenotypic analysis. The state of the art is a low cost and easy to set up 3D acquisition requiring only a background sheet, a reference object and a camera, compatible with field experiments in remote areas. We tested different software with CRC samples, and Agisoft and Blender were the most suitable software for generating high-quality 3D models and data analysis, respectively. We optimized the workflow by testing different numbers of images for 3D reconstruction and found that a minimum of 25 images per CRC can provide high quality 3D models. Up to ten traits, including 3D crown volumes, 3D crown surface, root density, surface-to-volume ratio, root numbers, root angle, crown diameter, cylinder soil volume, CRC compactness and root length can be extracted providing novel parameters for studying cassava storage roots. We applied this platform to partial-inbred cassava populations and demonstrated that our platform provides reliable 3D CRC modelling for phenotypic analysis, analysis of genetic variances and supporting breeding selection.

Validation of KASP-SNP markers in cassava germplasm for marker-assisted selection of increased carotenoid content and dry matter content.

Provitamin A biofortification and increased dry matter content are important breeding targets in cassava improvement programs worldwide. Biofortified varieties contribute to the alleviation of provitamin A deficiency, a leading cause of preventable blindness common among pre-school children and pregnant women in developing countries particularly Africa. Dry matter content is a major component of dry yield and thus underlies overall variety performance and acceptability by growers, processors, and consumers. Single nucleotide polymorphism (SNP) markers linked to these traits have recently been discovered through several genome-wide association studies but have not been deployed for routine marker-assisted selection (MAS). This is due to the lack of useful information on markers' performances in diverse genetic backgrounds. To overcome this bottleneck, technical and biological validation of the loci associated with increased carotenoid content and dry matter content were carried out using populations independent of the marker discovery population. In the present study, seven previously identified markers for these traits were converted to a robust set of uniplex allele-specific polymerase chain reaction (PCR) assays and validated in two independent pre-breeding and breeding populations. These assays were efficient in discriminating marker genotypic classes and had an average call rate greater than 98%. A high correlation was observed between the predicted and observed carotenoid content as inferred by root yellowness intensity in the breeding (r = 0.92) and pre-breeding (r = 0.95) populations. On the other hand, dry matter content-markers had moderately low predictive accuracy in both populations (r< 0.40) due to the more quantitative nature of the trait. This work confirmed the markers' effectiveness in multiple backgrounds, therefore, further strengthening their value in cassava biofortification to ensure nutritional security as well as dry matter content productivity. Our study provides a framework to guide future marker validation, thus leading to the more routine use of markers in MAS in cassava improvement programs.

Native and fermented waxy cassava starch as a novel gluten-free and clean label ingredient for baking and expanded product development.

Amylose-free and wild-type cassava starches were fermented for up to 30 days and oven- or sun-dried. The specific volume (ν) after baking was measured in native and fermented starches. The average ν (across treatments) for waxy starch was 3.5 times higher than that in wild-type starches (17.6 vs. 4.8 cm3 g-1). The best wild-type starch (obtained after fermentation and sun-drying) had considerably poorer breadmaking potential than native waxy cassava (8.4 vs. 16.4 cm3 g-1, respectively). The best results were generally obtained through the synergistic combination of fermentation (for about 10-14 days) and sun-drying. Fermentation reduced viscosities and the weight average molar mass led to denser macromolecules and increased branching degree, which are linked to a high loaf volume. The absence of amylose, however, was shown to be a main determinant as well. Native waxy starch (neutral in taste, gluten-free, and considerably less expensive than the current alternatives to cassava) could become a new ingredient for the formulation of clean label-baked or fried expanded products.

Development of waxy cassava with different Biological and physico-chemical characteristics of starches for industrial applications.

The quality of cassava starch, an important trait in cassava breeding programs, determines its applications in various industries. For example, development of waxy (having a low level of amylose) cassava is in demand. Amylose is synthesized by granule-bound starch synthase I (GBSSI) in plants, and therefore, down-regulation of GBSSI expression in cassava might lead to reduced amylose content. We produced 63 transgenic cassava plant lines that express hair-pin dsRNAs homologous to the cassava GBSSI conserved region under the control of the vascular-specific promoter p54/1.0 from cassava (p54/1.0::GBSSI-RNAi) or cauliflower mosaic virus (CaMV) 35S (35S::GBSSI-RNAi). After the screening storage roots and starch granules from field-grown plants with iodine staining, the waxy phenotype was discovered: p54/1.0::GBSSI-RNAi line A8 and 35S::GBSSI-RNAi lines B9, B10, and B23. Sodium dodecyl sulfate-polyacrylamide gel electrophoresis showed that there was no detectable GBSSI protein in the starch granules of plants with the waxy phenotype. Further, the amylose content of transgenic starches was significantly reduced (<5%) compared with the level in starch granules from the wild-type (about 25%). The inner structure of the waxy starch granules differed from that of the untransformed ones, as revealed by transmission electron microscopy analysis as well as morphological changes in the iodine-starch complex. Endothermic enthalpy was reduced in waxy cassava starches, according to differential scanning calorimeter analysis. Except B9, all waxy starches displayed the A-type X-ray diffraction pattern. Amylogram patterns of the waxy cassava starches were analyzed using a rapid viscosity analyzer and found to have increased values for clarity, peak viscosity, gel breakdown, and swelling index. Setback, consistency, and solubility were notably reduced. Therefore, waxy cassava with novel starch in its storage roots was produced using the biotechnological approach, promoting its industrial utilization.

Correlation of cooking time with water absorption and changes in relative density during boiling of cassava roots.

Consumers prefer cassava roots that cook quickly during boiling. Current methods to evaluate cooking time (CT) are slow and labour-intensive. This article describes improved protocols for assessing CT in roots. We evaluated CT in 36 genotypes monthly at 8-11 months after planting. CT showed differences for plant age at harvest and among genotypes. During boiling, roots absorbed water (WAB) and thus reduced their relative density (DEN). We classified three groups of genotypes with increasing CT (≤25 min, 25-40 min and >40 min), associated with decreasing WAB, respectively, 15.3 ± 3.1, 10.7 ± 1.7 and 4.9 ± 3.8% of initial root weight. A similar trend was observed for changes in DEN (46.3 ± 9.8, 54.5 ± 11.1 and 75.9 ± 6.9% of initial DEN, respectively). The highest correlations between WAB and DEN with CT (r 2 > 0.6) were found at 30-min boiling. These alternative protocols facilitate screening large numbers of cassava genotypes for CT.

Effect of Pruning Young Branches on Fruit and Seed Set in Cassava.

Flowering in cassava is closely linked with branching. Early-flowering genotypes branch low and abundantly. Although farmers prefer late flowering genotypes because of their erect plant architecture, their usefulness as progenitors in breeding is limited by their low seed production. In general, the first inflorescence aborts in cassava. Preventing this abortion would result in early production of seeds and make cassava breeding more efficient. The objective of this study was to assess if pruning young branches prevents the abortion of first inflorescences and promotes early fruit and seed set. Four genotypes with early, late, very late, and no flowering habits were grown under an extended photoperiod (EP) or normal dark night conditions (DN). Additional treatments included pruning young branches at the first or second flowering event and spraying (or not) benzyladenine (BA) after pruning. One genotype failed to flower and was not considered further. For the remaining genotypes, EP proved crucial to induce an earlier flowering, which is a pre-requisite for pruning. Total production of seeds in EP plots was 2,971 versus 150 in DN plots. For plants grown under EP, the average number of seeds per plant without pruning was 3.88, whereas those pruned produced 17.60 seeds per plant. Pruning at the first branching event led to higher number of seeds per plant (26.25) than pruning at the second flowering event (8.95). In general, applying BA was beneficial (38.52 and 13.98 seeds/plant with or without spraying it, respectively). The best combination of treatments was different for each genotype. Pruning young branches and applying BA in the first flowering event not only prevented the abortion of inflorescences but also induced the feminization of male flowers into hermaphrodite or female-only flowers. The procedures suggested from this study (combining EP, pruning young branches, and spraying BA), allowed the production of a high number of seeds from erect cassava genotypes in a short period. The implementation of these procedures will improve the breeding efficiency in cassava.

Selection mapping of loci for quantitative disease resistance in a diverse maize population.

The selection response of a complex maize population improved primarily for quantitative disease resistance to northern leaf blight (NLB) and secondarily for common rust resistance and agronomic phenotypes was investigated at the molecular genetic level. A tiered marker analysis with 151 simple sequence repeat (SSR) markers in 90 individuals of the population indicated that on average six alleles per locus were available for selection. An improved test statistic for selection mapping was developed, in which quantitative trait loci (QTL) are identified through the analysis of allele-frequency shifts at mapped multiallelic loci over generations of selection. After correcting for the multiple tests performed, 25 SSR loci showed evidence of selection. Many of the putatively selected loci were unlinked and dispersed across the genome, which was consistent with the diffuse distribution of previously published QTL for NLB resistance. Compelling evidence for selection was found on maize chromosome 8, where several putatively selected loci colocalized with published NLB QTL and a race-specific resistance gene. Analysis of F2 populations derived from the selection mapping population suggested that multiple linked loci in this chromosomal segment were, in part, responsible for the selection response for quantitative resistance to NLB.