Genetic variability of cassava progenies developed through introgression of cassava mosaic disease resistance into Ghanaian landraces

Availability of genetic diversity is important in selecting suitable cultivars for crop improvement. Progenies of crosses between 19 farmer-preferred cassava landraces and Cassava Mosaic Disease (CMD)-resistant cultivar TME 11 were evaluated and compared with their maternal parents to assess their genetic variability and genetic gains based on CMD tolerance, some growth variables and root yield. The combined analysis of variance for the traits showed significant genotypic differences amongst the progenies and the parents. Apart from a few instances, the differences in replications and years were also significant. Genetic variability components showed high heritability ranging from 92.3 to 99.6% and moderately high genetic gains of 54.1 to 99.6% for the parental cultivars. Apart from canopy spread and plant height, heritability and genetic gains of the progenies were slightly lower than the parents indicating high variability within families of the progenies. The phenotypic coefficient of variation was slightly greater than genotypic coefficient of variation indicating minimal influence of environment on the genotypes. For most of the characters evaluated, the mean performance of the progenies was higher than their corresponding parental lines. Although the genotypes differed significantly (P < 0.001) for all traits evaluated, cultivars such as Afebankye, Bosome nsia, Cedi bankye, 262 Debor, Kwadaso 25, Nkaakom 57 and Sisipe were found to constitute a pool of germplasm with adequate variability. The negative correlation between CMD incidence and other traits evaluated showed significant progress made in the breeding program.     

Diallel inheritance of relevant traits in cassava (Manihot esculenta Crantz) adapted to acid-soil savannas

There is a limited knowledge on the inheritance of traits with agronomic relevance in cassava. A diallel study among 10 parental clones was conducted in the acid-soils environment in the eastern savannas of Colombia. Thirty clones were obtained for each F1 cross. Each clone was represented by six plants, which were distributed in three replications at two locations. Therefore, the same 30 genotypes of each F1 cross were planted in the three replications at the two locations. Analysis of variance suggested significant effects for five of the six variables analyzed (harvest index, dry matter content, height of first branching, reaction to super elongation disease, and plant type scores). Fresh root yield showed strong genotype × environment interaction and differences between crosses reached statistical significance in only one of the two environments analyzed. General and specific combining ability effects and their interaction with the environment were significant for most of the variables as well. Results suggested that dominance plays an important role particularly in the cases of fresh root yield and harvest index.     

Correlated resistance of cassava to mosaic and bacterial blight diseases

Summary The two most serious diseases of cassava (Manihot esculenta Crantz) are cassava mosaic disease (CMD) and cassava bacterial blight (CBB) (Xanthomonas manihotis Starr). Clone 58308, derived from the third backcross of the interspecific cross of cassava (M. esculenta) x ceara rubber (M. glaziovii), showed a high level of resistance to both diseases. Crosses of 58308 with several other clones which varied from susceptible to moderately susceptible to both diseases gave progenies with a significant genotypic correlation between resistance to both diseases (r=0.90), apparently due to linkage. The heritabilities of resistance to the diseases were estimated at 50–70% for CMD and 25–65% for CBB. Resistance to both diseases is assumed to be polygenic. The correlated response to selection for CMD and for CBB was estimated.     

Agronomic performance and genotypic diversity for morphological traits among cassava genotypes in the guinea savannah ecology of Ghana

We evaluated the agronomic performance and diversity for morphological traits among a collection of local and exotic cassava genotypes in the Guinea Savannah agroecology of Ghana. One hundred and fifty cassava genotypes were assessed using 28 agro-morphological (18 qualitative and 10 quantitative) traits recorded from three months after planting until harvesting at 12 months after planting. There was significant genetic variability among the genotypes for most of the traits evaluated. High broad sense heritability was observed for most of the traits. Principal component analyses of the traits indicated that the first three principal component axes which accounted for more than 55% of the variation for 10 quantitative traits included storage root yield, number of leaves, harvest index, height at first branching, and plant height as the most important traits. A greater proportion of the phenotypic variances of plant height, height at branching, and harvest index were accounted for by genetic variances indicating that progress could made with selection. Pearson correlation analysis indicated that a number of traits were related to growth and yield of cassava in the Guinea Savanna ecology, making indirect selection for root yield possible. The study revealed ample genotypic variation in these morphological and agronomic traits to support selection for improved varieties.     

AMMI analysis of cassava response to contrasting environments: case study of genotype by environment effect on pests and diseases,root yield,and carotenoids content in Cameroon

Genotype by environment interaction remains a substantial issue in all breeding programs. Crop genotypes are generally developed in a central breeding location, but always require the evaluation of breeding products in different environments. This is particularly relevant in countries that have a wide range of climates. Eighteen cassava genotypes were evaluated in Cameroon in eight environments—varying in seasonal rainfall and temperature patterns and soil characteristics—over two cropping seasons. Soil nutrient content was analyzed and trials were established in a randomized complete block design in three replications. Response of genotypes to major cassava pests and diseases, yield and carotenoids content was evaluated. It was observed that four genotypes did not show cassava mosaic disease (CMD) symptoms irrespective of the environments. The local check had highest CMD incidence and severity across all environments. Average number of whitefly per plant across all environments was highest on TMS 96/0023. Average cassava green mite (CGM) infestation was low on all the genotypes. Fresh root yield of five genotypes ranged between 25 and 30 tons per ha for both years. Significant and positive correlation was found across locations between fresh root yield and soil K, P and Mg. AMMI analysis revealed highly significant differences among genotypes and environments and significant genotype?×?environment interaction for most of the estimated traits, indicating variability in genotypes performance with environment.     

The effect of cassava mosaic disease on the genetic diversity of cassava in Uganda

Summary Cassava (Manihot esculenta) is a tropical crop that is grown in Africa, Latin America and Southeast Asia. Cassava was introduced from Latin America into West and East Africa at two independent events. In Uganda a serious threat to cassava's survival is the cassava mosaic disease (CMD). Uganda has had two notable CMD epidemics since the introduction of cassava in the 1850s causing severe losses. SSR markers were used to study the effect of CMD on the genetic diversity in five agroecologies in Uganda with high and low incidence of CMD. Surprisingly, high gene diversity was detected. Most of the diversity was found within populations, while the diversity was very small among agroecological zones and the high and low CMD incidence areas. The high genetic diversity suggests a mechanism by which diversity is maintained by the active involvement of the Ugandan farmer in continuously testing and adopting new genotypes that will serve their diverse needs. However, in spite of the high genetic diversity we found a loss of rare alleles in areas with high CMD incidence. To study the effect of the introgression history on the gene pool the genetic differentiation between East and West Africa was also studied. Genetic similarities were found between the varieties in Uganda and Tanzania in East Africa and Ghana in West Africa. Thus, there is no evidence for a differentiation of the cassava gene pool into a western and an eastern genetic lineage. However, a possible difference in the genetic constitution of the introduced cassava into East and West Africa may have been diminished by germplasm movement.     

Standardisation of true cassava seed (TCS) programme with special emphasis on more homogeneous, CMD resistant progenies

The propagation of cassava through true seeds (sexual seeds) rather than by clones is a promising option due to its manifold advantages such as enhancing the multiplication rate, keeping the dreaded cassava mosaic disease (CMD) under check, longer seed viability, ease of storage and transport. The high genetic heterogeneity and consequent variation among seedlings is the major stumbling block in sexual propagation. In the present study, a CMD resistant exotic accession MNga-1 and a promising cultivar Ambakadan with profuse fruit setting, seed output and male sterility were identified to be promising parents for the TCS programme. The rate of sexual propagation could be more than 20-fold over the traditional clonal propagation. Seed treatment with 1% KNO₃ or 300 ppm GA promoted uniform seed germination and seedling vigour and reduced the transplanting period from 45 days after planting (DAS) to 30 DAS. Removal of taproots of seedlings while transplanting enhanced tuber development. Tuber yield of first clones (C₁) was significantly superior to that of the seedlings. The dry matter content and starch output of seedlings and first clones were comparable to that of the commercial varieties. Similarly, the HCN and cooking quality of seedlings and first clones were at acceptable levels. In the open pollinated (OP) progenies of the Ambakadan the CMD infection increased drastically due to secondary spread of the pathogen. The hybrid progenies of Ambakadan and the CMD resistant line MNga-1 revealed higher percentage of CMD free seedlings and first clonal progenies in the evaluation trials conducted at CTCRI Thiruvananthapuram and ARS, Peddapuram during 2001–2002 and 2002–2003. Nearly homogeneous hybrid population resistant to CMD could be obtained by systematic roguing at seedling and first clonal stage.     

Inheritance of useful traits in cassava grown in subhumid conditions

A diallel study among nine parental clones of cassava was conducted in the subhumid environment on the northern coast of Colombia. Analysis of variance suggested significant effects for the six variables analysed: fresh‐root yield, harvest index, root dry matter content, height of first branching, reaction to thrips and plant‐type scores. General and specific combining ability effects and their interaction with the environment were significant for most of the variables as well. Results suggested that dominance plays a particularly important role in the cases of fresh‐root yield and harvest index but had relatively little importance in the reaction to thrips, dry matter content or height of first branching. Specific breeding approaches are suggested for these traits, depending on the relative importance of additive or non‐additive effects in their inheritance. The correlations among different traits were also analysed and in several cases their magnitude reached statistical and biological significance.     

Morphological variation of starch granules in S<Subscript>1</Subscript> cassava progenies

The need to develop new cassava (Manihot esculenta Crantz) varieties with different starch characteristics has greatly motivated new research. The aim of this study was to evaluate morphological alterations in starch granules as well as genetic gains in segregating S₁ cassava populations. Starch from accessions (S₀): BGM0222, BGM1378 and BGM1662 and from their self-fertilized progenies (S₁): F0222, F1378 and F1662, with 78, 80 and 76 individuals, respectively, were evaluated for the following traits: area (ArStGr, µm²), perimeter (PrStGr, µm), length (LeStGr, µm) and number of starch granules (NuStGr). Moreover, some agronomic traits of the S₁ progenies were evaluated at harvest such as: plant height (PlaHei), shoot yield (ShoYi), root yield (RoYi), starch yield (StYi), and root dry matter content (DMC). With the selection of transgressive individuals in S₁ progenies, it is possible to obtain selection gains for values higher and lower than the average of the S₀ genitors, whereas selection gains can reach up to 16.58, 8.39, 7.87 and 34.18% for ArStGr, LeStGr, PrStGr, and NuStGr, respectively. Average size of the starch granule varied from 9.8 to 12.37 µm among individuals in an S₁ progeny. Positive and high correlations between the individuals were found for some morphological characteristics of the cassava starch granule but not strongly correlated with agronomic traits. Therefore, it is possible to select S₁ cassava individuals for agronomic characteristics without losing other important starch traits. Cassava-segregating populations allow the obtainment of individuals with different morphological characteristics as for starch granules, enabling obtainment of genetic gains with the selection.     

Association and Path Coefficients Analysis of Fresh Root Yield of High and Low Cyanide Cassava (Manihot esculenta Crantz) Genotypes in the Humid Tropics

Thirty-eight high- and low-cyanide cassava genotypes were examined in the 2004 and 2005 cropping seasons at Umudike, southeastern Nigeria using RCBD with three replications. Simple correlation associations and path coefficients analyses were computed among agronomic characters with the aim of understanding the interrelationships of these characters and the extent of their contribution to the root yield of cassava. There was a significant (P 2 0.05) positive correlation between root yield and number of roots plant-1 with correlation coefficients (r) of 0.57, average weight of root (r = 0.58) in high-cyanide cassava genotypes, while in low cyanide cassava genotypes, fresh root yield was highly significantly (P 2 0.01) correlated with average weight of root (r = 0.74). Path analysis showed that average weight of root had the highest direct effect in high (0.66) and in low cyanide (0.67) cassava genotypes. Also, the number of roots plant-1 in both types of cassava exerted a positive direct effect on fresh root yield, however, a minimum effect was obtained by number of leaves plant-1 (0.22) in high-cyanide cassava and number of stems plant-1 (-0.13) in lowcyanide cassava genotypes. Average weight of roots in high- and low-cyanide cassava genotypes had 98 and 89% direct effects, respectively, on yield of cassava. The results indicated the importance of average weight of roots and other characters such as number of roots plant-1, plant height, and number of leaves plant-1 in determining increased yield hence priority must be accorded it during selection aimed at increasing cassava root yield.     

Breeding cassava for resistance to cassava mosaic disease

Summary Cassava mosaic disease (CMD) is one of the most serious and widespread diseases throughout cassava growing areas in Africa, causing yield reductions of up to 90%. Early research on breeding of cassava (Manihot esculenta Crantz) for resistance to CMD in Africa is reviewed. Changes in population size and in activity of the white-fly vector to CMD (Bemisia tabaci Genn.) in relation to changes in environmental conditions such as amount and distribution of rainfall, light intensity and temperature are discussed in relation to screening for resistance to CMD. Over the past eight years, significant progress has been made at the International Institute of Tropical Agriculture (IITA). Resistance to CMD has been successfully incorporated into high yielding cultivars of acceptable quality. The CMD resistant material has been evaluated and many promising clones have been selected in various countries in tropical Africa and India. The resistance has been effective in those countries.     

Genotypic variation in abscisic acid content,carbon isotope ratio and their relationship with cassava growth and yield under moisture stress and irrigation

This study was carried out to assess genotypic variability in abscisic acid content, carbon isotope ratio, and their relationship to storage root yield and yield components in cassava under irrigation and moisture stress. The study involved 20 cassava genotypes arranged in randomized complete block design with three replications. Irrigation water was applied using a drip irrigation system with a discharge rate of approximately 5.33 L m^-2 hr^-1. Significant (P < 0.05) genotypic variability was observed for all physiological, growth, and yield traits assessed. Abscisic acid content was higher under stress than irrigation and negatively correlated with root yield (r = -0.45), harvest index (r = -0.43), and above-ground biomass yield (r = -0.20) indicating that it can be used as indirect selection criteria against unproductive genotypes. Carbon isotope ratio was significantly and positively correlated with above-ground biomass yield (r = 0.20) but not root yield (r = 0.09). Estimates of genotypic variability indicated high values for most of the growth and yield components but low heritability values for abscisic acid content, carbon isotope ratio, stomatal conductance, and root yield under stress conditions. However, higher estimates were recorded under irrigation and in the combined analysis. It was also found from this study that carbon isotope ratio influences above-ground biomass but not storage root yield under stress conditions. The results from this study provide useful information on the relationship between abscisic acid content, carbon isotope discrimination, and storage root yield in field-grown cassava.     

Adjustment for missing plants in cassava evaluation trials

Evaluations with a uniform plant stand are fundamental for efficient selections in cassava (Manihot esculenta Crantz) breeding. However, it is difficult to correct data of missing plants. The overall objective of this study was to analyze yield losses in agronomic trials and to develop a function that satisfactorily adjusts plot yields affected by a varying number of missing plants. Eight clones were initially evaluated in different locations for five consecutive years. For every genotype, mean plot yields decreased as the number of missing plants increased. Average losses ranged from 10.6 to 78.8% by removing from one to eight plants, respectively. Yields per plant increased significantly when more than four plants were removed due to a compensatory growth effect. Graphic analyses showed that the power function was the best explaining the relationship between fresh root yield and number of harvested plants. A model that properly adjusted yield for all varieties but one is proposed. Hopefully, this formula to adjust yields will help to improve the quality of cassava trials worldwide.     

Heterosis and Inbreeding Depression Estimation of Root and Shoot Characters in Rice (Oryza sativa L.)

The nature and extent of hybrid vigour and inbreeding depression were estimated from a diallel set of rice varieties. The seven parents and their 21 F₁s and 21 F₂s were grown in pots with four replications. Data were collected on four root and two shoot characters.
Maximum heterosis was recorded for root weight followed by roots/plant and root length. The inbreeding depression did not significantly affect the dry root weight in majority of the crosses. Heterosis for shoot weight was found to be associated with the heterosis for roots/plant and root weight.     

Genetic linkage map of cassava (Manihot esculenta Crantz) based on AFLP and SSR markers

To generate a genetic linkage map of cassava ( Manihot esculenta Crantz), 58 F₁ progenies from a cross between Rayong 90 (female) and Rayong 5 (male) were examined in amplification fragment length polymorphism (AFLP) and simple sequence repeat (SSR) analyses. A total of 469 polymorphic markers consisting of 378 AFLPs generated from 76 primer combinations and 91 SSRs were identified. These markers were analyzed using the joinmap ^® 3.0 package program to construct a genetic linkage map. A total of 33 linkage groups of a common map were constructed from 119 AFLPs and 18 SSRs, spanning 1095 cM with an average of 7.99 cM between markers. The genetic linkage map generated in this study will be useful for genetic studies in cassava particularly for the identification of genetic markers linked to traits of interest, although the complex cassava genome suggests that maybe a long term objective.     

Genome-wide selection in cassava

The main objective of this study was to estimate the selection accuracy and to predict the genetic gain in cassava breeding using genomic selection methodologies. We evaluated 358 cassava genotypes for the following traits: shoot weight (SW), fresh root yield (FRY), starch fraction amylose content (AC), dry matter content (DMC), and starch yield (S-Y). Genotyping was performed using 390 single nucleotide polymorphisms (SNPs), which were used as covariates in the random regression-best linear unbiased prediction model for genomic selection. The heritability values detected by markers for the SW, FRY, AC, DMC, and S-Y traits were 0.25, 0.25, 0.03, 0.20, and 0.26, respectively. Because the low heritability detected for AC, this trait was eliminated from further analysis. Using only the most informative SNPs (118, 92, 56, and 97 SNPs for SW, FRY, DMC, and S-Y, respectively) we observed higher selection accuracy which were 0.83, 0.76, 0.67, and 0.77, respectively to SW, FRY, DMC, and S-Y. With these levels of accuracy and considering a selection cycle reduced by half the time, the theoretical gains with genomic selection compared to phenotypic selection for DMC, FRY, and SW would be 39.42 %, 56.90 %, and 73.96 %, respectively. These results indicate that in the cassava, genomic selection can substantially speed up selection cycles, thereby increasing gains per unit time. Although there are high expectations for incorporating this strategy into breeding programs, we still need to validate the model for other traits and evaluate whether the selection accuracy can be improved using more SNPs.     

Development of superior cassava cultivars in Ghana by farmers and scientists: The process adopted, outcomes and contributions and changed roles of different stakeholders

Summary A participatory breeding programme involving farmers in two Ghanaian communities and scientists from CRI (Ghana) and NRI (UK) to develop superior cassava cultivars is described. Initial situation analyses of the communities indicated that cassava is increasing in importance both as a food and a cash crop. Most farmers utilised landraces of cassava; modern varieties were scarcely mentioned. Seeds of 16 half-sib families obtained from a crossing block in Nigeria at the International Institute of Tropical Agriculture were planted in a field in each community. During seedling and subsequent clonal generations, accessions selected either by farmers or scientists were retained to the next generation. This selection process has identified 29 superior accessions from amongst 1350 original seedlings. Farmers were relatively consistent in their selection from year to year and their selections corresponded with their stated criteria. Official variety release requires additional multilocational and inspection trials and postharvest assays but otherwise seems harmonious with a participatory breeding approach; our early involvement of farmers may facilitate early release, an important factor in cost-effectiveness. A stakeholder workshop confirmed the need for improved markets for cassava; surveys of current and potential markets have led to field trials with cassava processors. Adoption of a participatory approach, with farmers and scientists taking on new roles and decentralisation of activities, implies a concomitant transfer of influence and resources.     

Resistance profile of improved cassava germplasm to cassava mosaic disease in Nigeria

Cassava mosaic disease (CMD) caused by a group of begomoviruses and transmitted by whitefly vector is a serious disease in all the cassava-growing areas of Africa. Field evaluation with replication was conducted in 2003 and 2004 in three agroecologies in Nigeria to study the response of 40 cassava genotypes to CMD and to investigate genotype × environment (GE) interactions on their reactions to CMD, using the rank-sum classification and site regression analysis model. The 40 genotypes were separated into resistant (n = 17), moderately resistant (n = 6), moderately susceptible (n = 2) and susceptible (n = 15) groups. Environments, genotypes and GE interactions were all highly significant (P < 0.0001) for the virus disease contributing 9.5%, 71.36% and 19.14%, respectively to total variation. More than 40% of the genotypes were identified as resistant to the disease. Genotypes TMS 98/0581, TMS 99/3073, TMS 97/4763, TMS M98/0040, TMS 98/0505, TMS 97/0211, TMS 97/4769, TMS 99/2123, TMS M98/0068 and TMS 97/0162 were shown to have high resistance to CMD. The study also identified Umudike, in south-east Nigeria, as having high disease severity and the most appropriate site for CMD resistance screening of genotypes. Most of the genotypes exhibited stable resistance to CMD. The implication that the availability of these resistant genotypes as identified in this study could be a source of CMD resistance for further breeding is discussed.