Breeding Cultivars for Resistance to the African Sweetpotato Weevils, Cylas puncticollis and Cylas brunneus, in Uganda: A Review of the Current Progress.

In sub-Saharan Africa, sweetpotato weevils are the major pests of cultivated sweetpotato, causing estimated losses of between 60% and 100%, primarily during dry spells. The predominantly cryptic feeding behavior of Cylas spp. within their roots makes their control difficult, thus, host plant resistance is one of the most promising lines of protection against these pests. However, limited progress has been made in cultivar breeding for weevil resistance, partly due to the complex hexaploid genome of sweetpotato, which complicates conventional breeding, in addition to the limited number of genotypes with significant levels of resistance for use as sources of resistance. Pollen sterility, cross incompatibility, and poor seed set and germination in sweetpotato are also common challenges in improving weevil resistance. The accurate phenotyping of sweetpotato weevil resistance to enhance the efficiency of selection has been equally difficult. Genomics-assisted breeding, though in its infancy stages in sweetpotato, has a potential application in overcoming some of these barriers. However, it will require the development of more genomic infrastructure, particularly single-nucleotide polymorphism markers (SNPs) and robust next-generation sequencing platforms, together with relevant statistical procedures for analyses. With the recent advances in genomics, we anticipate that genomic breeding for sweetpotato weevil resistance will be expedited in the coming years. This review sheds light on Uganda's efforts, to date, to breed against the Cylas puncticollis (Boheman) and Cylas brunneus (Fabricius) species of African sweetpotato weevil.

Heterotic gains, transgressive segregation and fitness cost of sweetpotato weevil resistance expression in a partial diallel cross of sweetpotato.

Heterosis-exploiting breeding schemes are currently under consideration as a means of accelerating genetic gains in sweetpotato (Ipomoea batatas) breeding. This study was aimed at establishing heterotic gains, fitness costs and transgressive segregation associated with sweetpotato weevil (SPW) resistance in a partial diallel cross of sweetpotato. A total of 1896 clones were tested at two sites, for two seasons each in Uganda. Data on weevil severity (WED), weevil incidence (WI), storage root yield (SRY) and dry matter content (DM) were obtained. Best linear unbiased predictors (BLUPs) for each clone across environments were used to estimate heterotic gains and for regression analyses to establish relationships between key traits. In general, low mid-parent heterotic gains were detected with the highest favorable levels recorded for SRY (14.7%) and WED (- 7.9%). About 25% of the crosses exhibited desirable and significant mid-parent heterosis for weevil resistance. Over 16% of the clones displayed superior transgressive segregation, with the highest percentages recorded for SRY (21%) and WED (18%). A yield penalty of 10% was observed to be associated with SPW resistance whereas no decline in DM was detected in relation to the same. Chances of improving sweetpotato through exploiting heterosis in controlled crosses using parents of mostly similar background are somewhat minimal, as revealed by the low heterotic gains. The yield penalty detected due to SPW resistance suggests that a trade-off may be necessary between maximizing yields and developing weevil-resistant cultivars if the current needs for this crop are to be met in weevil-prone areas.

Targeting market segment needs with public-good crop breeding investments: A case study with potato and sweetpotato focused on poverty alleviation, nutrition and gender.

Crop breeding programs have often focused on the release of new varieties that target yield improvement to achieve food security and reduce poverty. While continued investments in this objective are justified, there is a need for breeding programs to be increasingly more demand-driven and responsive to the changing customer preferences and population dynamics. This paper analyses the responsiveness of global potato and sweetpotato breeding programs pursued by the International Potato Center (CIP) and its partners to three major development indicators: poverty, malnutrition and gender. The study followed a seed product market segmentation blueprint developed by the Excellence in Breeding platform (EiB) to identify, describe, and estimate the sizes of the market segments at subregional levels. We then estimated the potential poverty and nutrition impacts of investments in the respective market segments. Further, we employed the G+ tools involving multidisciplinary workshops to evaluate the gender-responsiveness of the breeding programs. Our analysis reveals that future investments in breeding programs will achieve greater impacts by developing varieties for market segments and pipelines that have more poor rural people, high stunting rates among children, anemia prevalence among women of reproductive age, and where there is high vitamin A deficiency. In addition, breeding strategies that reduce gender inequality and enhance appropriate change of gender roles (hence gender transformative) are also required.

Gender mainstreaming in sweetpotato breeding in Uganda: a case study.

Purpose: In Uganda, sweetpotato [Ipomoea batatas (L.) Lam] is typically a "woman's crop," grown, processed, stored and also mainly consumed by smallholder farmers for food and income. Farmers value sweetpotato for its early maturity, resilience to stresses, and minimal input requirements. However, productivity remains low despite the effort of breeding programs to introduce new varieties. Low uptake of new varieties is partly attributed to previous focus by breeders on agronomic traits and much less on quality traits and the diverse preferences of men and women in sweetpotato value chains. Method: To address this gap, breeders, food scientists, and social scientists (including gender specialists) systematically mainstreamed gender into the breeding program. This multidisciplinary approach, grounded in examining gender roles and their relationship with varietal and trait preferences, integrated important traits into product profiles. Results: Building on earlier efforts of participatory plant breeding and participatory varietal selection, new interventions showed subtle but important gender differences in preferences. For instance, in a study for the RTBFoods project, women prioritized mealiness, sweetness, firmness and non-fibrous boiled roots. These were further subjected to a rigorous gender analysis using the G+ product profile query tool. The breeding pipelines then incorporated these gender-responsive priority quality traits, prompting the development of standard operating procedures to phenotype these traits. Conclusion: Following an all-inclusive approach coupled with training of multidisciplinary teams involving food scientists, breeders, biochemists, gender specialists and social scientists, integration into participatory variety selection in Uganda enabled accentuation of women and men's trait preferences, contributing to clearer breeding targets. The research has positioned sweetpotato breeding to better respond to the varying needs and preferences of the users.

Household Processing Methods and Their Impact on Bioactive Compounds and Antioxidant Activities of Sweetpotato Genotypes of Varying Storage Root Flesh Colours.

Sweetpotato storage roots, peeled and unpeeled, of varying flesh colours (white, cream, yellow, pale orange, deep orange, and purple) were spectrophotometrically evaluated for their bioactive compounds and antioxidant activities. Roots were boiled, steamed, baked, fried, or microwaved. The unpeeled roots had relatively higher (p < 0.001) bioactive compounds and antioxidant activities than the peeled ones. All cooking methods increased phenolic compounds, flavonoids, and tannins in all genotypes. Significant losses of total carotenoids occurred with all cooking methods (ranging from 24.18 to 172.76 µg/g in raw sweetpotatoes vs. 10.06 to 118.17 µg/g in cooked ones; p < 0.001), except the deep-orange-fleshed genotype, in which frying slightly increased carotenoids from 269.81 to 304.74 µg/g. Microwaving retained 69% vitamin C in the cream-fleshed one, the highest among the cooking methods. Anthocyanins decreased with baking and frying in the purple-fleshed one but increased with other methods; microwaving being highest at 13.9% (17.43 mg/g). While the 2,2'-azino-bis-3-ethylbenzthiazoline-6-sulphonic acid antioxidant activity decreased with all cooking techniques in some genotypes, ferricyanide-reducing antioxidant potential increased. The retention of bioactive compounds in sweetpotato storage roots depends on the processing method. Thus, to obtain the most health benefits, consumers should use different cooking methods but retain the peels.

Combining ability and heritability analysis of sweetpotato weevil resistance, root yield, and dry matter content in sweetpotato.

Efficient breeding and selection of superior genotypes requires a comprehensive understanding of the genetics of traits. This study was aimed at establishing the general combining ability (GCA), specific combining ability (SCA), and heritability of sweetpotato weevil (Cylas spp.) resistance, storage root yield, and dry matter content in a sweetpotato multi-parental breeding population. A population of 1,896 F1 clones obtained from an 8 × 8 North Carolina II design cross was evaluated with its parents in the field at two sweetpotato weevil hotspots in Uganda, using an augmented row-column design. Clone roots were further evaluated in three rounds of a no-choice feeding laboratory bioassay. Significant GCA effects for parents and SCA effects for families were observed for most traits and all variance components were highly significant (p ≤ 0.001). Narrow-sense heritability estimates for weevil severity, storage root yield, and dry matter content were 0.35, 0.36, and 0.45, respectively. Parental genotypes with superior GCA for weevil resistance included "Mugande," NASPOT 5, "Dimbuka-bukulula," and "Wagabolige." On the other hand, families that displayed the highest levels of resistance to weevils included "Wagabolige" × NASPOT 10 O, NASPOT 5 × "Dimbuka-bukulula," "Mugande" × "Dimbuka-bukulula," and NASPOT 11 × NASPOT 7. The moderate levels of narrow-sense heritability observed for the traits, coupled with the significant GCA and SCA effects, suggest that there is potential for their improvement through conventional breeding via hybridization and progeny selection and advancement. Although selection for weevil resistance may, to some extent, be challenging for breeders, efforts could be boosted through applying genomics-assisted breeding. Superior parents and families identified through this study could be deployed in further research involving the genetic improvement of these traits.

Market Intelligence and Incentive-Based Trait Ranking for Plant Breeding: A Sweetpotato Pilot in Uganda.

Crop breeding programs must accelerate crop improvement, spur widespread adoption of new varieties and increase variety turnover they are to meet the diverse needs of their clients. More comprehensive quantitative approaches are needed to better inform breeding programs about the preferred traits among farmers and other actors. However, the ability of current breeding programs to meet the demands of their clients is limited by the lack of insights about value chain actor preference for individual or packages of traits. Ranking traits based on monetary incentives, rather than subjective values, represents a more comprehensive, consistent, and quantitative approach to inform breeding programs. We conducted a large pilot in Uganda to assess the implementation of a novel approach to trait ranking, using a uniquely large sample of diverse sweetpotato value chain actors. We found meaningful differences in trait ranking and heterogeneity among different actors using this approach. We also show our approach's effectiveness at uncovering unmet demand for root quality traits and at characterizing the substantial trait demand heterogeneity among value chain players. Implementing this approach more broadly for sweetpotato and other crops would increase the effectiveness of breeding programs to improve food security in developing countries.

The Potential of Sweetpotato as a Functional Food in Sub-Saharan Africa and Its Implications for Health: A Review.

Increasing urbanization in developing countries has resulted in busier lifestyles, accompanied by consumption of fast foods. The consequence is an increased prevalence in noncommunicable diseases (NCDs). Food-based approaches would be cheaper and more sustainable in reducing these NCDs compared to drugs, which may have side effects. Studies have suggested that consuming functional foods could potentially lower NCD risks. Sweetpotato is regarded as a functional food because it contains bioactive compounds. Recently, sweetpotato has gained attention in sub-Saharan Africa (SSA), but research has focused on its use in alleviating micronutrient deficiencies such as vitamin A deficiency, particularly the orange-fleshed variety of sweetpotato. Some studies conducted in other parts of the world have investigated sweetpotato as a functional food. There is a need to characterize the sweetpotato varieties in SSA and determine how processing affects their bioactive components. This review highlights some of the studies conducted in various parts of the world on the functionality of sweetpotato, its bioactive compounds, and how these are influenced by processing. In addition, the potential health benefits imparted by sweetpotato are expounded. The knowledge gaps that remain in these studies are also addressed, focusing on how they can direct sweetpotato research in SSA.

Discovery of a major QTL for root-knot nematode (Meloidogyne incognita) resistance in cultivated sweetpotato (Ipomoea batatas).

KEY MESSAGE: Utilizing a high-density integrated genetic linkage map of hexaploid sweetpotato, we discovered a major dominant QTL for root-knot nematode (RKN) resistance and modeled its effects. This discovery is useful for development of a modern sweetpotato breeding program that utilizes marker-assisted selection and genomic selection approaches for faster genetic gain of RKN resistance. The root-knot nematode [Meloidogyne incognita (Kofoid & White) Chitwood] (RKN) causes significant storage root quality reduction and yields losses in cultivated sweetpotato [Ipomoea batatas (L.) Lam.]. In this study, resistance to RKN was examined in a mapping population consisting of 244 progenies derived from a cross (TB) between 'Tanzania,' a predominant African landrace cultivar with resistance to RKN, and 'Beauregard,' an RKN susceptible major cultivar in the USA. We performed quantitative trait loci (QTL) analysis using a random-effect QTL mapping model on the TB genetic map. An RKN bioassay incorporating potted cuttings of each genotype was conducted in the greenhouse and replicated five times over a period of 10 weeks. For each replication, each genotype was inoculated with ca. 20,000 RKN eggs, and root-knot galls were counted ~62 days after inoculation. Resistance to RKN in the progeny was highly skewed toward the resistant parent, exhibiting medium to high levels of resistance. We identified one major QTL on linkage group 7, dominant in nature, which explained 58.3% of the phenotypic variation in RKN counts. This work represents a significant step forward in our understanding of the genetic architecture of RKN resistance and sets the stage for future utilization of genomics-assisted breeding in sweetpotato breeding programs.

Development of a food product profile for boiled and steamed sweetpotato in Uganda for effective breeding.

This study sought to understand user preferences of raw, boiled and steamed sweetpotato, a staple food in Uganda. A sequential methodology involving state of knowledge review, gendered food mapping, processing diagnosis and consumer testing was used in Lira and Kamwenge districts. Preferred raw sweetpotato characteristics were large roots (≥ 3 cm diameter) with a sweet taste, smooth skin and hard texture, while mealiness, sweet taste and good sweetpotato smell were important attributes for boiled sweetpotato. Processors, mostly women, highlighted ease of peeling and sappiness of raw roots. There were gender differences in quality characteristic preferences and perceived importance. The released variety, NASPOT 8, had the highest overall liking in Kamwenge and was well liked in Lira. Penalty analysis of consumer data showed that sweetness and firmness were key drivers of overall liking. The results will support breeding programmes in meeting specific end-user product profiles, selection criteria and uptake of new varieties.

Quantitative trait loci and differential gene expression analyses reveal the genetic basis for negatively associated ß-carotene and starch content in hexaploid sweetpotato [Ipomoea batatas (L.) Lam.].

KEY MESSAGE: ß-Carotene content in sweetpotato is associated with the Orange and phytoene synthase genes; due to physical linkage of phytoene synthase with sucrose synthase, ß-carotene and starch content are negatively correlated. In populations depending on sweetpotato for food security, starch is an important source of calories, while ß-carotene is an important source of provitamin A. The negative association between the two traits contributes to the low nutritional quality of sweetpotato consumed, especially in sub-Saharan Africa. Using a biparental mapping population of 315 F1 progeny generated from a cross between an orange-fleshed and a non-orange-fleshed sweetpotato variety, we identified two major quantitative trait loci (QTL) on linkage group (LG) three (LG3) and twelve (LG12) affecting starch, ß-carotene, and their correlated traits, dry matter and flesh color. Analysis of parental haplotypes indicated that these two regions acted pleiotropically to reduce starch content and increase ß-carotene in genotypes carrying the orange-fleshed parental haplotype at the LG3 locus. Phytoene synthase and sucrose synthase, the rate-limiting and linked genes located within the QTL on LG3 involved in the carotenoid and starch biosynthesis, respectively, were differentially expressed in Beauregard versus Tanzania storage roots. The Orange gene, the molecular switch for chromoplast biogenesis, located within the QTL on LG12 while not differentially expressed was expressed in developing roots of the parental genotypes. We conclude that these two QTL regions act together in a cis and trans manner to inhibit starch biosynthesis in amyloplasts and enhance chromoplast biogenesis, carotenoid biosynthesis, and accumulation in orange-fleshed sweetpotato. Understanding the genetic basis of this negative association between starch and ß-carotene will inform future sweetpotato breeding strategies targeting sweetpotato for food and nutritional security.

Genetic analysis and association of simple sequence repeat markers with storage root yield, dry matter, starch and ß-carotene content in sweetpotato.

Molecular markers are needed for enhancing the development of elite sweetpotato (Ipomoea batatas (L.) Lam) cultivars with a wide range of commercially important traits in sub-Saharan Africa. This study was conducted to estimate the heritability and determine trait correlations of storage root yield, dry matter, starch and ß-carotene content in a cross between 'New Kawogo' × 'Beauregard'. The study was also conducted to identify simple sequence repeat (SSR) markers associated with these traits. A total of 287 progeny and the parents were evaluated for two seasons at three sites in Uganda and genotyped with 250 SSR markers. Broad sense heritability (H2) for storage root yield, dry matter, starch and ß-carotene content were 0.24, 0.68, 0.70 and 0.90, respectively. Storage root ß-carotene content was negatively correlated with dry matter (r = -0.59, P < 0.001) and starch (r = -0.93, P < 0.001) content, while storage root yield was positively correlated with dry matter (r = 0.57, P = 0.029) and starch (r = 0.41, P = 0.008) content. Through logistic regression, a total of 12, 4, 6 and 8 SSR markers were associated with storage root yield, dry matter, starch and ß-carotene content, respectively. The SSR markers used in this study may be useful for quantitative trait loci analysis and selection for these traits in future.

Segregation of Hydroxycinnamic Acid Esters Mediating Sweetpotato Weevil Resistance in Storage Roots of Sweetpotato.

Resistance to sweetpotato weevils (Cylas spp.) has been identified in several sweetpotato (Ipomoea batatas) landraces from East Africa and shown to be conferred by hydroxycinnamic acids that occur on the surface of storage roots. The segregation of resistance in this crop is unknown and could be monitored using these chemical traits as markers for resistance in F1 offspring from breeding programs. For the first time in a segregating population, we quantified the plant chemicals that confer resistance and evaluated levels of insect colonization of the same progeny in field and laboratory studies. We used a bi-parental mapping population of 287 progenies from a cross between I. batatas 'New Kawogo,' a weevil resistant Ugandan landrace and I. batatas 'Beauregard' a North American orange-fleshed and weevil susceptible cultivar. The progenies were evaluated for resistance to sweetpotato weevil, Cylas puncticollis at three field locations that varied climatically and across two seasons to determine how environment and location influenced resistance. To augment our field open-choice resistance screening, each clone was also evaluated in a no choice experiment with weevils reared in the laboratory. Chemical analysis was used to determine whether differences in resistance to weevils were associated with plant compounds previously identified as conferring resistance. We established linkage between field and laboratory resistance to Cylas spp. and sweetpotato root chemistry. The data also showed that resistance in sweetpotato was mediated by root chemicals in most but not all cases. Multi-location trials especially from Serere data provided evidence that the hydroxycinnamic acid esters are produced constitutively within the plants in different clonal genotypes and that the ecological interaction of these chemicals in sweetpotato with weevils confers resistance. Our data suggest that these chemical traits are controlled quantitatively and that ultimately a knowledge of the genetics of resistance will facilitate management of these traits, enhance our understanding of the mechanistic basis of resistance and speed the development of new sweetpotato varieties with resistance to sweetpotato weevil.