Biology subjects

Rabbi, I. Y.

Publications and source records attributed to Rabbi, I. Y..

13 recordsLinked to original sources

bioRxiv2026

Breeding cassava for intercropping with cowpea: monoculture selection captures most intercrop selection gain, but targeted testing remains necessary

Intercropping dominates smallholder cassava production in sub-Saharan Africa, yet cassava breeding programs evaluate genotypes exclusively under monoculture. Despite consistently reported system-level yield advantages, cassava yield is reduced by 17 to 51% under intercropping, indicating a need to reduce this competitive disadvantage through breeding. However, the quantitative-genetic foundations of intercrop breeding remain uncharacterized for tropical root crop systems. We hypothesized that monoculture selection would capture most, but not all, genetic merit for intercropping and that a limited tester set would be sufficient if general mixing ability predominated. We evaluated 120 cassava clones previously selected under monoculture in IITA advanced yield trials, testing them under monoculture and in intercrop with two contrasting cowpea varieties across two years at Ibadan, Nigeria. Spatial mixed models quantified genetic variation, genotype by cropping system interaction, cross system genetic relationships, realized selection gain, mixing ability, tester effects, and land equivalent ratio. Intercropping reduced cassava fresh root yield by 19%, but total land equivalent ratios exceeded 1.0 for all clones, confirming a system-level land use advantage. Cassava performance under intercropping was heritable, with estimates of 0.50 to 0.75, and genotype by cropping system interaction was not significant. Genetic correlations between monoculture and intercrop performance were high and approached unity (rg = 0.92 to 0.99), and selection efficiency was 26 to 44% at 10% intensity, confirming that high genetic correlation does not guarantee effective indirect selection. Monoculture selection captured approximately two-thirds of direct intercrop gain. General mixing ability dominated, specific mixing ability was negligible, and producer effects explained 20 to 47% of intercrop variance. The two architecturally and phenologically contrasting cowpea varieties had limited influence on cassava rankings. Here, we show for the first time that cassava breeding for intercropping can retain monoculture selection during early stages while adding two representative cowpea testers at the advanced trial stage. This staged strategy aligns cassava breeding with diversified smallholder systems without creating a separate pipeline.

plant biology↗

bioRxiv2026

Repeatability and Heritability of UAV-Derived Canopy Traits in a Cassava Breeding Population Using Time-Series Data from Two Consecutive Growing Seasons

Cassava is a major staple crop in tropical regions, particularly in Sub-Saharan Africa, yet its productivity remains constrained by genetic and agronomic limitations. A major bottleneck in cassava breeding is the difficulty of accurately phenotyping agronomic traits under field conditions using conventional, labor-intensive methods. Here, we evaluated the potential of uncrewed aerial vehicle (UAV)-based phenotyping to quantify canopy growth traits and assess their genetic relevance under realistic field conditions. For this, multi-temporal UAV imagery was collected over two growing seasons (2018-2019 and 2019-2020) in a panel of 46 cassava genotypes planted in fields of the International Institute for Tropical Agriculture (IITA), Nigeria. Canopy height, canopy volume, and their relative growth rates (RGRh and RGRv) were extracted at the plot-level, and their seasonal dynamics and canopy-yield relationships were further assessed across developmental stages and environmental conditions. Repeatability (R) and broad-sense heritability (H2) were estimated using a linear mixed model (LMM) that partitioned genetic, genotype-by-year, and residual variance components, enabling the evaluation of both measurement reliability and genetic signal. Overall, UAV-derived growth dynamics were found to exhibit comparable patterns across genotypes, reflecting shared seasonal growth trajectories, while canopy-yield relationships varied with developmental stage and environmental conditions. In terms of genetic metrics, R was high for all UAV-derived traits (R = 0.68-0.69), indicating reliable genotype-level assessment across replicates and seasons. In contrast, H2 differed substantially among traits. Canopy volume (H2 = 0.64) and canopy height (H2 = 0.58) exhibited moderate-to-high heritability, reflecting strong genotype effects and comparatively moderate genotype-by-year interactions. However, their relative growth rates showed near-zero H2 values, driven primarily by genotype-by-year interaction, indicating a dominant environmental influence. These results demonstrate that UAV-derived canopy height and volume provide a consistent basis for genetic differentiation of cassava genotypes across environments, supporting their use in selection, whereas growth-rate traits are better suited for characterizing growth plasticity and genotype-by-environment interactions.

plant biology↗

bioRxiv2026

Introgression from the wild relative Manihot glaziovii on cassava (M. esculenta) chromosome 1 exhibits segregation distortion and no direct effect on dry matter

The cassava (Manihot esculenta) genome has two large introgressions from its wild relative M. glaziovii on chromosomes 1 and 4 that originate from historical hybridization efforts. The 10 Mbp chromosome 1 introgression has been increasing in frequency in African breeding populations due to its statistical association with higher dry matter content and root number. However, the region also exhibits suppressed recombination, hindering breeders ability to combine favorable glaziovii alleles with the cultivated esculenta background. Since homozygous introgressed lines are rarely selected for advanced trials, dominance effects have not been well-characterized. To analyze the effects of the introgression with higher resolution, we generated a population of over 5000 seedlings from crosses between heterozygous introgressed parents and screened for recombinants using ten KASP markers tagging glaziovii-specific alleles. An optimized subset of 453 lines was then selected and evaluated over two years for yield and vigor traits. Unlike previous studies, composite interval mapping and mixed linear models showed no significant associations between glaziovii alleles and dry matter content or root number. Small, opposing effects on clonal vigor were observed at different ends of the introgression. The region showed significant segregation distortion and enrichment of putative deleterious alleles. Genome alignment of M. esculenta and M. glaziovii assemblies did not show any major structural variants in the introgression region, suggesting that suppressed recombination is likely driven by sequence-level divergence rather than structural rearrangements. These results indicate that the glaziovii introgression does not directly contribute to dry matter, supporting the need for recombination and purging of the glaziovii introgression to aid cassava improvement. Plain language summaryA large chromosome segment from a wild relative of cassava is an important structural aspect in the cassava genome. Since the chromosome segment tends to be inherited as one block, its effects on cassava traits were not well resolved. Through genetic mapping at higher resolution, we identified that the wild segment impacts early vigor and does not appear to impact dry yield, as was previously thought. While there are no major structural differences between the wild and cultivated chromosome segments, their overall divergence seems to suppress the wild chromosome segment from pairing with the cultivated chromosome segment during reproduction. In the apparent absence of any major benefits from the wild segment, removing it from the breeding population may be beneficial. Core ideasO_LIA set of glaziovii allele-specific markers were designed to track the chromosome 1 introgression haplotype. C_LIO_LISegregation distortion suggests the presence of recessive deleterious or lethal alleles in the introgression. C_LIO_LIIncreased recombination is needed to purge deleterious alleles enriched in introgression region. C_LIO_LIThe glaziovii introgression was associated with slightly lower vigor rating and stem diameter. C_LIO_LIThe effects of the previously-identified glaziovii DM QTL were not detected in this population. C_LI

genetics↗

bioRxiv2026

Genetic basis of Cassava (Manihot esculenta Crantz) plant architecture and its relevance for selection of farmer-preferred varieties

Plant architecture, the spatial configuration of stems, branches, leaves, and inflorescences underpins essential physiological functions such as light capture, assimilate partitioning, flowering, and ultimately, yield. In cassava (Manihot esculenta), architectural traits such us plant height, branching level, and plant shape are agronomically important yet remain underexploited in breeding. Here, a large-scale analysis was conducted using phenotypic and genomic data from more than 14,000 cassava accessions evaluated across 34 field locations in Nigeria between 2010 and 2021, encompassing the national breeding programs of the National Root Crops Research Institute and the International Institute of Tropical Agriculture. The study aimed to dissect the genetic architecture, environmental stability, and breeding relevance of four key traits: plant full height, height to first branching, the branching level number (BranchlevelNum) and plant shape. Phenotypic analyses across breeding stages revealed consistent variation in plant height, branching height, and branching intensity, reflecting the cumulative effects of selection and evaluation across environments. Broad-sense heritability estimates ranged from 0.41 to 0.72, with BranchlevelNum and Cylindrical shape exhibiting strong genetic control and weak correlations with yield components, indicating their suitability for independent improvement. Genome-wide association analyses identified significant loci associated with BranchlevelNum, including a major region on chromosome 2 and an additional locus on chromosome 13, collectively explaining approximately 11% of the phenotypic variance. Candidate genes within these regions included regulators of meristem activity and hormone-related pathways, supporting a developmental basis for branching variation. Genomic prediction accuracy for BranchlevelNum reached 0.44, comparable to values reported for key agronomic traits in cassava. These results demonstrate that branching-related architectural traits are genetically tractable, largely independent of yield, and amenable to genomic selection. The findings support the integration of BranchlevelNum and plant shape into ideotype-driven breeding frameworks aimed at improving flowering efficiency, canopy structure, and field performance in cassava. Author SummaryCassava is a major food crop, and its plant shape plays an important role in how easily it can be grown, harvested, and improved through breeding. Traits such as plant height, branching, and canopy form affect flowering, seed production, and field management, yet they have received much less attention than yield or disease resistance. In this study, we examined plant architecture using field and genetic data from more than 14,000 cassava plants grown across Nigeria over twelve years. We focused on key traits describing plant height, branching level, and overall plant shape. We found that branching level is strongly controlled by genetics, remains stable across environments, and can be predicted accurately using genomic data. We also identified specific regions of the cassava genome linked to branching behavior. Our findings show that plant architecture can be improved using modern breeding tools without compromising yield. Incorporating branching traits into breeding programs can help develop cassava varieties that flower more reliably and perform better in farmers fields.

genetics↗

bioRxiv2024

Evaluating a Cassava Crop Growth Model by Optimizing Genotypic-Specific Parameters Using Multi-environment Trial Breeding Data

Cassava (Manihot esculenta Crantz) is a critical food security crop for sub-Saharan Africa. Efforts to improve cassava through breeding have expanded over the past decade. At the same time, crop growth models (CGM) are becoming common place in breeding efforts to expand the inference of evaluations of breeding germplasm to environments that have not been tested and to prepare for breeding for adaptation to future climates. We parameterized a CGM, the CROPGRO-MANIHOT-Cassava model in the DSSAT family of models, using data on 67 clones from the International Institute of Tropical Agriculture cassava breeding program evaluated from 2017 to 2020 and over eight locations in Nigeria using trial and error parameter adjustments and the General Likelihood Uncertainty Estimation method. Our objectives were to assess the feasibility of this large-scale calibration in the context of a cassava breeding program and to identify systematic biases of the model. For each cultivar we calculated the Pearson correlation between model prediction and observation across the environments, as well as root mean squared error and d statistics. As a result of calibration, the correlation coefficient increased from -0.03 to +0.08, the RMSE dropped from 21 t ha-1 to 5 t ha-1 while d increased from 0.23 to 0.44. We found that the model underestimated root yield in dry environments (low precipitation and high temperature) and overestimated root yield in wet environments (high precipitation and low temperature). Our experience suggests both that CGM calibration could become a routine component of the cassava breeding data analysis cycle and that there are opportunities for model improvement.

plant biology↗

bioRxiv2023

Carbon usage in yellow-fleshed Manihot esculenta storage roots shifts from starch biosynthesis to cell wall and raffinose biosynthesis via the myo-inositol pathway

Cassava is a crucial staple crop for smallholder farmers in tropical Asia and Sub-Saharan Africa. Although high yield remains the top priority for farmers, the significance of nutritional values has increased in cassava breeding programs. A notable negative correlation between provitamin A and starch accumulation poses a significant challenge for breeding efforts. The negative correlation between starch and carotenoid levels in conventional and genetically modified cassava plants implies the absence of a direct genomic connection between the two traits. The competition among various carbon pathways seems to account for this relationship. In this study, we conducted a thorough analysis of 49 African cassava genotypes with varying levels of starch and provitamin A. Our goal was to identify factors contributing to differential starch accumulation. With the carotenoid levels of the varieties considered as a confounding effect on starch production, we found that yellow and white-fleshed storage roots did not differ significantly in most measured components of starch or de novo fatty acid biosynthesis. However, genes and metabolites associated with myo-inositol synthesis and cell wall component production were substantially enriched in high provitamin A genotypes. These results indicate that yellow-fleshed cultivars, in comparison to their white-fleshed counterparts, direct more carbon towards the synthesis of raffinose and cell wall components, a finding that is supported by a significant rise in the starch-free residue to total dry yield ratio in yellow storage roots versus white storage roots. Our findings enhance comprehension of the biosynthesis of starch and carotenoids in the storage roots of cassava.

plant biology↗

bioRxiv2022

Validation of SNP markers for marker-assisted selection of genotypes with increased carotenoid and dry matter contents in cassava

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 in developing countries. Dry matter content is a major component of dry yield and thus underlies overall variety performance and acceptability by growers, processors, and consumers. SNP markers linked to these traits have recently been discovered through several genome-wide association studies but have not been deployed for marker-assisted selection (MAS). Assessment of marker performance in diverse genetic backgrounds is an important step towards their deployment for routine MAS. In the present study, seven previously identified markers for these traits were converted to a robust set of uniplex allele-specific 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 marker effectiveness in multiple backgrounds, therefore, further strengthening its value in cassava biofortification to ensure nutritional security as well as dry matter content productivity. Our study provides a framework to guide future marker development, thus leading to more routine use of markers in MAS in cassava improvement programs.

genomics↗

bioRxiv2021

Low-cost, handheld near-infrared spectroscopy for root dry matter content prediction in cassava

Over 800 million people across the tropics rely on cassava as a major source of calories. While the root dry matter content (RDMC) of this starchy root crop is important for both producers and consumers, characterization of RDMC by traditional methods is time-consuming and laborious for breeding programs. Alternate phenotyping methods have been proposed but lack the accuracy, cost, or speed ultimately needed for cassava breeding programs. For this reason, we investigated the use of a low-cost, handheld NIR spectrometer for field-based RDMC prediction in cassava. Oven-dried measurements of RDMC were paired with 21,044 scans of roots of 376 diverse clones from 10 field trials in Nigeria and grouped into training and test sets based on cross-validation schemes relevant to plant breeding programs. Mean partial least squares regression model performance ranged from R2p = 0.62 - 0.89 for within-trial predictions, which is within the range achieved with laboratory-grade spectrometers in previous studies. Relative to other factors, model performance was highly impacted by the inclusion of samples from the same environment in both the training and test sets. Random forest variable importance analysis of root spectra revealed increased importance in a region previously identified as predictive of water content in plants (~950 - 990 nm). With appropriate model calibration, the tested spectrometer will allow for field-based collection of spectral data with a smartphone for accurate RDMC prediction and potentially other quality traits, a step that could be easily integrated into existing harvesting workflows of cassava breeding programs. CORE IDEASO_LIA low-cost, handheld near-infrared spectrometer was tested for phenotyping of cassava roots C_LIO_LIPlant breeding-relevant cross-validation schemes were used for predictions C_LIO_LIHigh prediction accuracies were achieved for cassava root dry matter content C_LIO_LIA spectral region predictive of plant water content was identified as important C_LI

plant biology↗

bioRxiv2020

Genetic architecture and gene mapping of cyanide in cassava (Manihot esculenta Crantz.)

Cassava is a root crop originating from South America and a major staple crop in the Tropics, including marginal environments. In this study, we focused on South American and African cassava germplasm and investigated the genetic architecture of Hydrogen Cyanide (HCN), a major component of tuber quality. HCN is a plant defense component against herbivory but also toxic for human consumption. We genotyped 3,354 landraces and modern breeding lines originating from 26 Brazilian states and 1,389 individuals were phenotypically characterized across multi-year trials for HCN. All plant material was subjected to high density genotyping using Genotyping-by-sequencing (GBS). We performed genome wide association mapping (GWAS) to characterize the genetic architecture and gene mapping of HCN. Field experiment revealed strong broad and narrow-sense trait heritability (0.82 and 0.41 respectively). Two major loci were identified, encoding for an ATPase and a MATE protein and contributing up to 7% and 30% of the cyanide concentration in roots, respectively. We developed diagnostic markers for breeding applications, validated trait architecture consistency in African germplasm and investigated further evidence for domestication of sweet and bitter cassava. Fine genomic loci characterization indicate; (i) a major role played by vacuolar transporter in regulating HCN content, (ii) co-domestication of sweet and bitter cassava major alleles to be geographical zone dependant, and (ii) major loci allele for high cyanide cassava in Manihot esculenta Crantz seems to originate from its ancestor, M. esculenta ssp. flabellifolia. Taken together these findings expand insights on cyanide in cassava and its glycosylated derivatives in plants. One-sentence summaryIdentification of an intracellular transporter gene and its allelic variation allow to point out cultivars with up to 30 percent decrease in cassava root cyanide content, toxic for human consumption.

genetics↗

bioRxiv2020

Genome-wide association analysis reveals new insights into the genetic architecture of defensive, agro-morphological and quality-related traits in cassava

Cassava (Manihot esculenta) is one of the most important starchy root crops in the tropics due to its adaptation to marginal environments. Genetic progress in this clonally propagated crop can be accelerated through the discovery of markers and candidate genes that could be used in cassava breeding programs. We carried out a genome-wide association study (GWAS) using a panel of 5,310 clones developed at the International Institute of Tropical Agriculture - Nigeria. The population was genotyped at more than 100,000 SNP markers via genotyping-by-sequencing (GBS). Genomic regions underlying genetic variation for 14 traits classified broadly into four categories: biotic stress (cassava mosaic disease and cassava green mite severity); quality (dry matter content and carotenoid content) and plant agronomy (harvest index and plant type). We also included several agro-morphological traits related to leaves, stems and roots with high heritability. In total, 41 significant associations were uncovered. While some of the identified loci matched with those previously reported, we present additional association signals for the traits. We provide a catalogue of favourable alleles at the most significant SNP for each trait-locus combination and candidate genes occurring within the GWAS hits. These resources provide a foundation for the development of markers that could be used in cassava breeding programs and candidate genes for functional validation.

genetics↗

bioRxiv2019

Improving root characterisation for genomic prediction in cassava

Cassava is widely cultivated due to its high drought tolerance and high carbohydrate-containing storage roots. The lack of uniformity and irregular shape of storage roots within and between genotypes poses significant constraints on harvesting and post-harvest processing. Routine assessment of storage root size and shape in breeding plots relies on visual scores.\n\nHere, we phenotyped the Genetic gain and offspring (C1) populations from the International Institute of Tropical Agriculture (IITA) breeding program for root shape and size-related traits using image analysis of storage root photographs taken in the field.\n\nIn our study, using univariate genome-wide association analysis, we detected for most shape and size related traits, significant QTL regions located on chromosomes 1 and 12. The QTL region on chromosome 12 has been previously associated, using IITA breeding populations, to cassava mosaic disease (CMD) resistance.\n\nBecause the uniformity in size and shape of cassava roots is an important breeding goal, we calculated the standard deviation of individual root measurements per clone. The use of standard deviation measurements allowed the identification of new significant QTL for Perimeter, Feret and Aspect Ratio on chromosomes 6, 9 and 16. Using genomic prediction cross validation, the accuracies of root size and shape-related traits were lower than those previously reported for dry matter content (DM) and cassava mosaic virus resistance (CMD). Predictive accuracies of the mean values of root size and shape image-extracted traits were mostly higher than yield trait prediction accuracies in the C1 population. This study aimed to evaluate the feasibility of the image phenotyping protocol and to assess the use of genome-wide analyses for size and shape image-extracted traits. The methodology described here and the results obtained in this study are promising and open up the opportunity to apply high-throughput methods in cassava.

genomics↗

bioRxiv2017

Prospects for genomic selection in cassava breeding

Cassava (Manihot esculenta Crantz) is a clonally propagated staple food crop in the tropics. Genomic selection (GS) reduces selection cycle times by the prediction of breeding value for selection of unevaluated lines based on genome-wide marker data. GS has been implemented at three breeding programs in sub-Saharan Africa. Initial studies provided promising estimates of predictive abilities in single populations using standard prediction models and scenarios. In the present study we expand on previous analyses by assessing the accuracy of seven prediction models for seven traits in three prediction scenarios: (1) cross-validation within each population, (2) cross-population prediction and (3) cross-generation prediction. We also evaluated the impact of increasing training population size by phenotyping progenies selected either at random or using a genetic algorithm. Cross-validation results were mostly consistent across breeding programs, with non-additive models like RKHS predicting an average of 10% more accurately. Accuracy was generally associated with heritability. Cross-population prediction accuracy was generally low (mean 0.18 across traits and models) but prediction of cassava mosaic disease severity increased up to 57% in one Nigerian population, when combining data from another related population. Accuracy across-generation was poorer than within (cross-validation) as expected, but indicated that accuracy should be sufficient for rapid-cycling GS on several traits. Selection of prediction model made some difference across generations, but increasing training population (TP) size was more important. In some cases, using a genetic algorithm, selecting one third of progeny could achieve accuracy equivalent to phenotyping all progeny. Based on the datasets analyzed in this study, it was apparent that the size of a training population (TP) has a significant impact on prediction accuracy for most traits. We are still in the early stages of GS in this crop, but results are promising, at least for some traits. The TPs need to continue to grow and quality phenotyping is more critical than ever. General guidelines for successful GS are emerging. Phenotyping can be done on fewer individuals, cleverly selected, making for trials that are more focused on the quality of the data collected.\n\nAbbreviations

genetics↗

bioRxiv2017

Genome-wide association mapping of correlated traits in cassava: dry matter and total carotenoid content

Cassava (Manihot esculenta (L.) Crantz) is a starchy root crop cultivated in the tropics for fresh consumption and commercial processing. Dry matter content and micronutrient density, particularly of provitamin A - traits that are negatively correlated - are among the primary selection objectives in cassava breeding. This study aimed at identifying genetic markers associated with these traits and uncovering the potential underlying cause of their negative correlation - whether linkage and/or pleiotropy. A genome-wide association mapping using 672 clones genotyped at 72,279 SNP loci was carried out. Root yellowness was used indirectly to assess variation in carotenoid content. Two major loci for root yellowness was identified on chromosome 1 at positions 24.1 and 30.5 Mbp. A single locus for dry matter content that co-located with the 24.1 Mbp peak for carotenoid content was identified. Haplotypes at these loci explained a large proportion of the phenotypic variability. Evidence of mega-base-scale linkage disequilibrium around the major loci of the two traits and detection of the major dry matter locus in independent analysis for the white- and yellow-root subpopulations suggests that physical linkage rather that pleiotropy is more likely to be the cause of the negative correlation between the target traits. Moreover, candidate genes for carotenoid (phytoene synthase) and starch biosynthesis (UDP-glucose pyrophosphorylase and sucrose synthase) occurred in the vicinity of the identified locus at 24.1 Mbp. These findings elucidate on the genetic architecture of carotenoids and dry matter in cassava and provides an opportunity to accelerate genetic improvement of these traits.\n\nCORE IDEASO_LICassava, a starchy root crop, is a major source of dietary calories in the tropics.\nC_LIO_LIMost varieties consumed are poor in micronutrients, including pro-vitamin A.\nC_LIO_LIThese two traits are governed by few major loci on chromosome one.\nC_LIO_LIGenetic linkage, rather than pleiotropy, is the most likely cause of their negative correlation.\nC_LI

genetics↗