Abstract
BACKGROUND
Vitamin A deficiency (VAD) is widespread in sub‐Saharan Africa (SSA). Unlike in developed countries, where the main source of vitamin A comes from meat, the diet of poor populations in SSA is largely plant based. It is thus important to identify local / popular plants with higher vitamin A content for combating VAD. Banana (including plantains) is an important staple food crop in this region. The identification and promotion of vitamin A‐rich banana cultivars could contribute significantly to the alleviation of VAD in areas heavily dependent on the crop. We assessed pro‐vitamin A carotenoid (pVACs) content in the fruit pulp of 48 local plantains from eastern Democratic Republic of Congo, to identify cultivars that could help reduce VAD, especially among young children and women of reproductive age.
RESULTS
Mean pVACs content varied from 175–1756 μg/100 gfw in ripe fruits. Significant increases (P < 0.001) in total pVACs content occurred after ripening in all cultivars except ‘UCG II’. Retinol activity equivalents (RAE) in ripe fruits ranged from 12–113 μg/100 gfw. Fifteen plantain cultivars, including ‘Adili II’, ‘Nzirabahima’, ‘Mayayi’, ‘Buembe’, and ‘Sanza Tatu’ (associated with RAE values of 44 μg/100 gfw and above) can be considered as good sources of pVACs. Modest consumption (250 or 500 gfw) of the fruit pulp of the five best plantain cultivars at ripening stage 5 meets between 39–71% and 44–81% of vitamin A dietary reference intake (DRI) respectively, for children below 5 years old and women of reproductive age.
CONCLUSION
The 15 best plantain cultivars (especially the top 5) could potentially be introduced / promoted as alternative sources of pro‐vitamin A in banana‐dependent communities, and help to reduce cases of VAD substantially. © 2019 The Authors. Journal of The Science of Food and Agriculture published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.
Keywords: children below 5 years, dietary reference intake (DRI), pro vitamin A, retinol activity equivalents (RAE), vitamin A deficiency, women of child‐bearing age
INTRODUCTION
Vitamin A (retinol) and its derivatives (retinal and retinoic acid) are essential for cell differentiation, proliferation, and signaling, and they thus play an important role in vision, reproduction, maintenance of epithelial tissue, and immune functions.1, 2, 3
Vitamin A deficiency (VAD) is a major public health problem in many developing countries due to an over‐reliance on starchy staples and processed foods that are typically deficient in vitamin A, with preschool children and pregnant women being the most vulnerable groups.4, 5 Vitamin A deficiency causes serious chronic diseases such as night blindness, keratomalacia (i.e. dryness with ulceration and perforation of the cornea), loss of vision, bronchopulmonary dysplasia, growth retardation, shortening and thickening of bones, atrophy of the testes, fetal reabsorption, immunodeficiency, and increased morbidity and mortality from infectious diseases.1, 2, 3, 6, 7 It has been estimated that VAD affects over 190 million children below 5 years of age, especially in Africa and Southeast Asia.4, 5 Over the period 1995 to 2005, up to 1.2 million deaths among children aged 1 to 4 years worldwide were linked to VAD, amongst other factors.5, 8 In 2007, the prevalence of VAD in the Democratic Republic of Congo (DR Congo) and Burundi was established at 42.2% (25.2 out of 59.8 million) and 42.6% (3.38 out of 7.94 million) respectively, and has been classified as a ‘severe public health problem’ by the United Nations Standing Committee on Nutrition.9
Humans cannot synthesize vitamin A, and so rely on their diet for its supply, which can be from animal origins in the form of preformed vitamin A (retinol) and / or plant origin in the form of pro‐vitamin A carotenoids (pVACs).10, 11, 12 Preformed vitamin A from animal sources (retinol) is more easily absorbed and metabolized than the pVACs from plant foods, which have to be converted into retinol in the human body to be used.13, 14, 15 But foods of animal origin are not always available for millions of people across the world.16 White‐fleshed roots and tubers (e.g. cassava, sweet potato, taro, and yam), banana, and rice dominate people's diets in most households in developing countries, and these are often low in pVACs. The bioavailability of pVACs from different plant sources also varies considerably.17, 18 Furthermore, the structure of pVACs is paramount; for instance, the retinol activity equivalent (RAE) obtained upon the intake of a single unit of all‐trans β‐carotene (t‐BC) is twice that of all‐trans α‐carotene.14 It is thus important not only to identify and incorporate plant sources that have high levels of pVACs but also those that naturally contain a high proportion of t‐BC.
Commonly known plant sources that are rich in pVACs include yellow and orange fruits such as mangoes and papayas, orange‐fleshed sweet potatoes, and dark‐green leafy vegetables.16, 19 However, consumption is often limited in resource‐poor communities, due to the high perishability (coupled to a lack of storage facilities), seasonality, limited accessibility, and high cost of the fruits and / or vegetables. For example, a 2007 survey carried out in North Kivu province, eastern DR Congo reported that only 55% of children aged 6 to 35 months had consumed foods rich in vitamin A in the previous 24 h, which is lower than the national average of 66%.20
Approaches that have been used to meet vitamin A dietary reference intake (DRI) include supplementation (i.e. administering concentrated doses of micronutrients to at‐risk populations) and food fortification (i.e. adding micronutrients to processed food).21, 22 Although supplementation and food fortification were found to be effective in several studies,5 both have generally proven difficult to implement in developing countries,23 as they are mainly urban based and often fail to reach the rural communities most in need. These communities most often have a limited access to medical facilities and cannot afford fortified foods. With plant sources accounting for over 80% of vitamin A intake in low‐income countries,10, 24 the promotion of a diverse diet that includes vitamin A‐rich local food crops is now thought to be a more sustainable strategy. In the case of DR Congo, 58% of breastfed children 6–23 months of age, and 38% of those not breastfed, consume diets with three or less food groups, thus having low dietary diversity.20 Similarly, a study in Butembo, North Kivu found that 48% of children aged 2–5 years had poorly diversified diets.25
In large parts of highland East and Central Africa, the food system is banana‐based, with daily consumption levels of highland banana (AAA‐EAHB) fruits in countries such as Uganda, Rwanda, and Burundi varying between 0.3 and 1.6 kg per person.26, 27, 28, 29 The carotenoid content in the fruit pulp of banana has been shown to vary between and within cultivars with geographical location, time of harvest, and ripening stage30, 31, 32, 33, 34, 35, 36 (Table 1). The identification and promotion of vitamin A‐rich cultivars is considered a feasible approach to help alleviate VAD. The widespread promotion of vitamin A‐rich banana (including plantain) cultivars has, for example, been reported to lead potentially to a reduction of 9.6–17.1% in the burden of illnesses due to VAD in three African countries: Ghana, Rwanda, and Uganda.30
Table 1.
Range of pro‐vitamin A content for various banana cultivars and genome groups, and other crops as reported in the literature. Data are reported as listed in the respective publications. For Mbabazi (2015),36 values per gdw were converted into values per 100 gfw using moisture data
| Reference | Crop/ banana cultivar (group) | Range of total pro‐vitamin A content (μg/100 gfw) |
|---|---|---|
| Englberger et al. (2003)34 | Micronesian bananas | 515 to 6360 μg/100 gfw |
| Englberger et al. (2003)35 | Southeast Asia banana cultivars | 300 to 4960 μg/100 gfw |
| Englberger et al. (2006)37 | Micronesian Fe'i cultivar ‘Utimwas’ (boiled) | 9000 μg/100 gfw |
| Boiled Pacific plantain (AAB) cultivar ‘Mangat en Seipahn’ | 8207 μg/100 gfw | |
| Davey et al. (2009)30, 38 | 171 Musa cultivars covering essentially all genome groups | 0 to 3457 μg/100 gfw |
| Ekesa et al. (2013)39 | Selected cooking (2) and plantain (2) cultivars from eastern DR Congo | 76–182 μg/100 gfw |
| Mbabazi (2015)36 | Dessert bananas (Sukari Ndizi and Gros Michel) | 6.3 to 14.4 μg/gdw (∼270 to 620 μg/100 gfw) |
| East African Highland banana (AAA‐EA) cultivars | 9.6 to 19.9 μg/gdw (∼413 to 856 μg/100 gfw) | |
| Plantain cultivar (AAB) ‘Gonja Nakatansese’ | 37 μg/gdw (∼1591 μg/100 gfw) | |
| Transgenic ‘Sukari Ndizi’ with genes enhanced for pVACs production | 12.6 to 17.3 μg/gdw (∼542 to 744 μg/100 gfw) | |
| Fungo et al. (2010)40 | Various East African Highland Banana (AAA‐EA) cultivars | 146.4–513.7 μg/100 gfw |
| ‘Sukari ndizi’ | 50.6 μg/100 gfw | |
| Institute of Medicine (US) Panel on Micronutrients (2002)14 | Carrots | 11 427 to 14 693 μg/100 gfw |
| Tomato | 3454 μg/100 gfw | |
| Spinach | 9890 μg/100 gfw | |
| Green cabbage | 139 μg/100 gfw | |
| Englberger et al. (2006)41, 42 | Nine Pandanus cultivars of the Kiribati republic | 154 to 3602 μg/100 gfw |
| 13 Pandanus cultivars of the Republic of the Marshall Island | 260 to 3130 μg/100 gfw | |
| van Jaarsveld et al. (2005)43 | Orange‐fleshed sweet potato | 8329–10 699 μg/100 gfw |
| Welsch et al. (2010)44 | White‐fleshed cassava | 60 to 150 μg/100 gfw |
| Tan (1989)45 | Unrefined/crude palm oil | RAE 15 times higher than carrots, and 300 times that of tomatoes |
In eastern DR Congo, plantain (AAB genome) and green cooking banana (AAA genome, East African highland banana – EAHB) have been reported as the second most important source of calories, after cassava.39 Recent studies have reported plantains and EAHB to be potential sources of pVACs.32, 38, 39, 46, 47 However, the above studies on pVACs in banana have focused on a limited number of EAHB and plantain cultivars. Screening of a wider range of plantain cultivars was deemed necessary to identify those cultivars that could meet or exceed the DRIs of vitamin A for susceptible / vulnerable groups. This study therefore assessed, through C30‐RP‐HPLC analysis, the pVACs content at ripening stages 1 and 5 for 48 plantain cultivars grown in different parts of the North Kivu and Ituri provinces of eastern DR Congo, which is also part of a secondary center of banana diversity.
MATERIALS AND METHODS
Study region and plantain cultivars
The eastern part of the DR Congo covers a wide variety of agro‐ecologies that support a wide diversity of bananas, including plantains.48 It is characterized by two rainy seasons (mid‐August to mid‐January and mid‐February to mid‐July) and two short dry seasons (mid‐January to mid‐February and mid‐July to mid‐August), and a mean precipitation varying between 1000 and 3170 mm. Altitude ranges from 700 to 5119 m above sea level (a.s.l.) (Mt Rwenzori top), and mean annual temperature varies from as low as 15 °C (high altitude) to as high as 31 °C (low altitude).28, 49, 50 This region also has a high variability in its soils, including volcanic soils, alluvial soils, naturally weathered deep and humus‐rich rock soils.28
This study was conducted on 48 different local plantain cultivars (AAB), previously collected through a Musa germplasm survey covering three districts in North Kivu province and five in Ituri province of eastern DR Congo (Table 2 51). It is estimated that the plantains used in this study represent about 30% of the known plantain diversity across DR Congo. Table 2 provides details of the cultivar types and their relative abundance at the sites of origin.
Table 2.
Plantain cultivars assessed for agronomic performance and pro‐vitamin A carotenoid content in eastern Democratic Republic of Congo
| Cultivar name | Typea | Site of origin | Relative abundanceb | Citationc |
|---|---|---|---|---|
| Adili I | F | Ituri | * | A |
| Adili II | F | Ituri | * | A |
| Agbindolo | F | Ituri | ** | A |
| Akange | F | Ituri | * | A |
| Akobanzi | FH | Ituri | * | A |
| Akoto membo | FH | Ituri | * | A |
| Akoto monama | FH | Ituri | * | A |
| Alongo | FH | Ituri | * | A |
| Apakumo | FH | Ituri | *** | A |
| Ayaya | F | Ituri | * | A |
| Bakpulu | FH | Ituri | * | A |
| Bubu | FH | Ituri | * | A |
| Buembe | F | North Kivu | * | A, B |
| Kingulungulu | FH | North Kivu | * | A, B |
| Kirisirya plantain | F | North Kivu | * | A, B |
| Kothina | FH | North Kivu | ** | A, B |
| Kothina I | FH | North Kivu | ** | A |
| Kothina II | FH | North Kivu | ** | A |
| Kothina III | FH | North Kivu | * | A |
| Mabilanga | FH | Ituri | * | A |
| Makaka | H | Ituri | * | A |
| Makpelekese | FH | Ituri | * | A |
| Mangondi | FH | Ituri | *** | A |
| Mangondi I | FH | Ituri | * | A |
| Mangondi II | FH | Ituri | * | A |
| Manzenzele | F | Ituri | * | A |
| Mayayi | F | Ituri | * | A |
| Musilongo | FH | North Kivu | ** | A, B |
| Ndonge | F | Ituri | * | A |
| Ngobia II‐1 | F | Ituri | * | A |
| Ngobia makelekele | FH | Ituri | * | A, B |
| Nguma | F | North Kivu | *** | A, B |
| Nguma II | F | North Kivu | ** | A |
| Nguma III | F | North Kivu | *** | A |
| Nzirabahima | F | North Kivu | * | A, B |
| Plantain masunga | F | North Kivu | * | A |
| Plantain grand format I | FH | North Kivu | * | A |
| Plantain grand format II | FH | North Kivu | * | A |
| Sanza moya | H | North Kivu/Ituri | * | A, B |
| Sanza tatu | H | North Kivu | * | A, B |
| UCG II | FH | Ituri | * | A |
| UCG III | FH | North Kivu | * | A |
| UCG IV | F | North Kivu | * | A |
| UCG VIII | F | Ituri | * | A |
| Vuhembe | F | North Kivu | * | A, B |
| Vuhetera | F | North Kivu | * | A, B |
| Vuhindi | F | North Kivu/Ituri | ** | A, B |
| Vulambya | F | North Kivu | * | A, B |
Field experimental design and data collection
Field experiments were established in September 2010 at the Catholic University of Graben's (UCG) regional Musa germplasm collection at Butembo (1815 m a.s.l., 00.11786° N, 29.2587° E) in North Kivu province, for assessment of plantain cultivar yield and pro‐vitamin A content. Butembo has sandy clay soils with pH 5.7, 5.1% organic carbon (OC), 0.26% nitrogen, 10.6 mg kg−1 of available phosphorus and 1.14 cmol(+) kg−1 of potassium.51 The site had a mean annual rainfall of 1036 mm and a mean annual temperature of 19 °C.
For each of the 48 plantain cultivars, 15 plants were grown in three replications of five plants each, at a spacing of 2 m × 3 m. Banana production was entirely rain fed. No fertilizers and chemicals (for pest or disease control) were applied, and the fields were regularly mulched with dry grass.
The 48 plantain cultivars were observed during the period 2010–2015, over two cropping cycles, i.e. the plant and first ratoon crop. Data collected for this study included bunch weight (kg) and the time to harvest (days). Time to harvest for the plant and first ratoon crop is, respectively, time from planting to harvest and time from harvest of the plant crop to harvest of the first ratoon crop.
Average annual yield (tons/ha/year) was based upon the two crop cycles and was calculated from bunch weight and the time to harvest using the formula described by Gaidashova et al. (2008)52 as below: Average annual yield = [(bunch weight (kg)/number of days to harvest] × 365 × 1666.7)/1000; where 365 = number of days in a year; 1666.7 = plant density/ha, and 1000 = conversion factor from kg to tons.
Sample collection and preparation
The samples for pVACs analysis were collected from the Butembo field collection. Here, three physiologically mature disease‐free plant crop bunches (when fruits were deep green, full, and rounded; i.e. ripening stage 153) were selected per cultivar and harvested over a period spread out between 2012 and 2013. Two middle hands (second and third hand from the top of the bunch) per bunch were subsequently sampled (giving a total of six hands per cultivar) and used for carotenoid analysis. The fresh fruit samples were packed in perforated cardboard boxes and transported under ambient temperature (20 °C) to the National Agricultural Research Laboratory, Kawanda, Kampala, Uganda, within 24 h of harvest for further processing prior to analysis. Fruit ripeness was assessed based on peel color as described by Dadzie and Orchard (1997)53 and Stover and Simmonds (1987):54 1 = green; 2 = green with trace of yellow; 3 = more green than yellow; 4 = more yellow than green; 5 = only green tips remaining; 6 = all yellow; 7 = yellow flecked with brown. In this study, fruits were sampled for pVACs assessment at ripening stages 1 (unripe) and 5 (ripe) as they reflect the extreme range of plantain pulp consumption. Plantains are mainly consumed (fried or boiled) at ripening stage 5. Nevertheless, unripe plantain is also often fried or boiled. At ripening stage 1, three fingers were randomly detached from each of the six banana hands per cultivar, cleaned, hand peeled, and cut into pieces. Fruit pulp pieces from the two hands collected from the same bunch were combined into a sub‐sample, resulting in three subsamples per cultivar. Each sub‐sample was placed in a labeled, zip‐locked bag from which air was manually removed, and frozen at −80 °C. The remaining fingers on the six selected hands per cultivar were left to ripen naturally in a well aerated room. Sub‐sampling of fruit pulp at ripening stage 5 was done using the same procedure applied during stage 1. Once both stages were sampled, samples were weighed and lyophilized for 72 h, after which they were re‐weighed, re‐packaged in labeled zip‐locked bags, and kept frozen at −80°C until transport to Belgium for nutrient analyses. Half of each sub‐sample was transported on dry ice, using a courier service, to the Laboratory of Fruit Breeding and Biotechnology, Department of Bio‐systems at the Catholic University of Leuven (KUleuven), Belgium for pVACs analysis.
Extraction and analysis of carotenoids
Carotenoid extraction and analysis were carried out in triplicate according to procedures developed specifically for the analysis of banana tissues as described in Ekesa et al. (2013b).39 One hundred milligram aliquots of powdered lyophilized fruit pulp were homogenized for 30 s at maximum speed in a ‘Fast Prep’ reciprocal shaker in 400 mL of ice‐cooled extraction solvent (Thomas Scientific Inc., Swedesboro, New Jersey, USA). The extraction solvents consisted of tetrahydrofuran : methanol (THF : MeOH), 1:1 (v/v), containing 0.25% butylated hydroxytoluene (BHT) and 2% insoluble polyvinylpolypyrolidine (PVPP) to complex flavonoids and other secondary metabolites that can interfere with analyses.31, 55 Following centrifugation (16,873 xg for 20 min at 48 °C), the supernatant was transferred to a fresh micro‐centrifuge tube (Heathrow Scientific LLC, Vernon Hills, Illinois, USA) and the pellet was re‐extracted twice with 400 mL of THF : MeOH, 1:1 (v/v), containing 0.25% BHT without PVPP. Supernatants were collected and combined, and 8‐apo β‐carotenal at a final concentration of 0.002 mg/mL was added as an internal standard. The combined supernatants were analyzed directly by reversed phase – high performance liquid chromatography (RP‐HPLC) using a Waters Alliance, 2690 Separations System fitted with an auto‐sampler, thermostat at 88 °C, a pulse dampener and a 996 UV–visible photodiode array detector (Waters, Milford, MA, USA). The entire system was controlled and the data were collected and integrated using the Millennium 4.0 software package. A 150 mm ‐ 4.6 mm, YMC C30 3 μm particle size HPLC column (Achrom, Zulte, Belgium) was employed, using a 24 min linear gradient from 2 to 50% tert‐butyl methyl ether (t‐BME) in MeOH at 1.0 mL/min, followed by a 1 min linear gradient to 95% t‐BME in MeOH, which was maintained for 3 min to elute the remaining compounds. The column was then re‐equilibrated for 5 min under starting conditions of 2% t‐BME in MeOH, before the next injection. Peaks were quantified at 450 nm using a freshly prepared standard curve of all‐trans‐ β‐carotene and 8‐apo‐carotenal in extraction solvent, and identified on the basis of their characteristic absorption spectra and retention times relative to known standards.39, 55
Pro‐vitamin A carotenoids data conversions
Results from the pVACs analysis were obtained as nmol/g dry weight (gdw) and converted to nmol/gfw [fresh matter weight = dry matter weight/ (100/ (100‐moisture %))] and finally to μg/100 gfw using the conversion factor of 0.01863.56 Fresh matter weight was measured after peeling the fruit, and dry matter weight was measured after lyophilization. To determine the relative vitamin A nutritional content of samples, total pVACs content were first converted to all‐trans‐β‐carotene equivalents (t‐BCEs) by adding up all the specific carotenoids, which included all‐trans‐β‐carotene (t‐BC), all‐trans‐a‐carotene (t‐AC) and their cis‐isomers (c‐BC, c‐AC). As t‐AC has half of the retinol activity of t‐BC, and c‐BC and c‐AC each have 0.53 times the retinol activity of t‐BC, the following formula [t‐BCE = 0.5 t‐AC + t‐BC + 0.53 (c‐BC + c‐AC)] was used.55, 57, 58
On average, the consumption of 12 μg of t‐BC from fruits and vegetables yields 1 μg of retinol (Institute of Medicine, 2001), therefore, t‐BCE values were converted into retinol activity equivalents (RAE) assuming that 1/12th of the total t‐BCEs ingested are taken up by the body.59
The DRIs of vitamin A for children below 5 years old and of women of reproductive age (15–49 years) equal 400 μg RAE and 700 μg RAE, respectively.60 The RAE values (in μg/100 gfw) of the studied plantain fruits were compared to the above daily vitamin A DRIs when varying amounts of fruit pulp were consumed. Consumption of 250 g (approximately 1–2 fingers) of cooked plantain fruit by a child below 5 years and 500 g (approximately 3–4 fingers) by a woman of reproductive age has been reported to be within normal consumption levels in banana growing and consuming regions of Africa.61 However, the above amounts may not always be attained across the various banana‐based production systems in Africa. Therefore, DRI values were also calculated for smaller fruit pulp quantities, varying between 100–250 g for children below 5 years and 100–500 g for women of reproductive age. Taking into account the exceptionally high fruit pulp consumption levels of adults in the east and central African highland regions, higher DRI values can also be derived from values calculated for 100, 250, and 500 g of fruit pulp for women of reproductive age. The percentage DRI for the various cultivars and fruit ripening stages and for different fruit pulp quantities consumed was then computed as: (RAE of cultivar [μg/X gfw] × 100)/DRI, with DRI = 400 μg RAE for children below 5 years and 700 μg RAE for women of reproductive age.
Statistical analysis
The analysis of variance (ANOVA) for yield (tons/ha/year), individual pVACs, total pVACs content, total t‐BCEs, RAE, and DRIs for children below 5 years and women of reproductive age across the 48 plantain cultivars and the two ripening stages were determined using GenStat v. 12 statistical software.62 The Least Significant Difference (LSD) function in GenStat v.12 was used to compare means at 5% significance level. In addition, GenStat v. 12 was also used to carry out regression analysis between on one hand plant growth cycle duration and bunch weights with pVACs content on the other hand.
RESULTS AND DISCUSSION
Provitamin A carotenoids content and RAE of plantains
The pVACs identified in the plantain cultivars under study were t‐BC, t‐AC, cis‐AC and cis‐BC (Table 3, Figs. 1 and 2). Trace amounts of all‐trans‐lutein were also recorded although this compound has no vitamin A activity. Similar pVACs profiles were reported for different banana cultivars by Davey et al. (2009)30, 31, 38 and Ekesa et al. (2013, 2015).33, 39, 47 At ripening stage 1, the plantain cultivars had a similar pVACs profile containing mainly t‐BC with lower levels of t‐AC, except ‘Akange’, which contained only t‐BC, ‘Kirisirya Plantain’ (primarily lutein) and ‘Plantain Grand Format’, which consisted primarily of t‐AC (Fig. 1). At ripening stage 5, the plantain cultivars had a similar pVACs profile with higher proportions of t‐BC (47–70%) followed by t‐AC (23–43%) and only trace amounts of cis‐BC, cis‐AC, and lutein (Fig. 2). This is similar to results from Davey et al. (2009)30, 38 and Ekesa et al. (2015)33 who reported a larger proportion of t‐BC (61 to 69%) compared to t‐AC (30–38%) in some plantain cultivars; t‐AC has only 50% of the RAE of t‐BC and thus the relative proportions of t‐AC and t‐BC affect the overall vitamin A value in foods.57, 58 The proportion of t‐BC relative to other pVACs is therefore a good indicator of the quality of vitamin A supplied by a given food. Studies on other crops such as maize and wheat have reported a much lower t‐BC proportion of 10% to 20% of the total carotenoid content.38 Thus the quality of the pVACs in the 48 plantains is good compared to some other plant pVACs sources.
Table 3.
Mean content of individual and total pro‐vitamin A carotenoids (pVACs) and all‐trans‐β‐carotene equivalents (t‐BCE) in μg/100 gfw in fruit pulp of 48 plantain (AAB) cultivars at ripening stages 1 and 5. Values are means of three individual samples per ripening stage
| Cultivar name | Ripening stage | t‐AC | t‐BC | Cis‐AC | Cis‐BC | Lutein | Total‐pVACs | t‐BCE |
|---|---|---|---|---|---|---|---|---|
| Adili I | 1 | 109.2 | 183.6 | 7.3 | 0.0 | 7.7 | 300.1 | 242.1 |
| 5 | 283.1 | 311.4 | 15.9 | 0.0 | 48.9 | 610.4 | 461.4 | |
| Adili II | 1 | 16.5 | 20.9 | 0.0 | 0.0 | 0.0 | 37.4 | 29.2 |
| 5 | 740.5 | 958.3 | 9.9 | 47.4 | 152.1 | 1756.1 | 1358.9 | |
| Agbindolo | 1 | 71.4 | 162.7 | 1.2 | 5.7 | 2.6 | 241.0 | 202.0 |
| 5 | 162.9 | 336.3 | 26.7 | 17.8 | 23.4 | 543.7 | 441.4 | |
| Akange | 1 | 0.0 | 4.0 | 0.0 | 0.0 | 0.0 | 4.0 | 4.0 |
| 5 | 46.9 | 127.3 | 7.4 | 0.0 | 8.7 | 181.6 | 154.7 | |
| Akobanzi | 1 | 4.8 | 7.6 | 0.0 | 0.0 | 0.0 | 12.4 | 10.0 |
| 5 | 47.2 | 118.1 | 9.2 | 0.0 | 11.6 | 174.5 | 146.5 | |
| Akoto membo | 1 | 1.5 | 10.2 | 0.0 | 0.0 | 0.0 | 11.7 | 10.2 |
| 5 | 334.8 | 502.7 | 42.2 | 0.0 | 24.9 | 879.7 | 692.5 | |
| Akoto monama | 1 | 24.9 | 30.3 | 0.0 | 0.0 | 2.3 | 55.2 | 42.8 |
| 5 | 272.5 | 389.7 | 14.7 | 0.0 | 10.9 | 676.9 | 533.8 | |
| Alongo | 1 | 8.9 | 15.4 | 0.0 | 0.0 | 0.0 | 24.3 | 19.9 |
| 5 | 154.3 | 360.9 | 17.1 | 0.0 | 14.0 | 532.3 | 447.1 | |
| Apakumo | 1 | 41.6 | 108.2 | 0.0 | 0.0 | 0.0 | 149.8 | 129.0 |
| 5 | 124.0 | 289.7 | 0.0 | 12.4 | 0.0 | 426.1 | 358.2 | |
| Ayaya | 1 | 8.3 | 20.1 | 0.0 | 0.0 | 0.0 | 28.4 | 24.3 |
| 5 | 163.5 | 266.1 | 12.2 | 0.0 | 21.0 | 441.8 | 354.3 | |
| Bakpulu | 1 | 10.4 | 26.3 | 0.0 | 0.0 | 0.0 | 36.7 | 31.5 |
| 5 | 93.0 | 238.2 | 20.5 | 0.0 | 12.9 | 351.7 | 295.6 | |
| Bubu | 1 | 17.5 | 47.1 | 0.0 | 0.0 | 0.0 | 64.6 | 55.8 |
| 5 | 238.5 | 509.3 | 0.0 | 37.7 | 24.7 | 785.5 | 648.5 | |
| Buembe | 1 | 31.2 | 80.4 | 0.0 | 2.6 | 0.0 | 114.2 | 97.4 |
| 5 | 311.7 | 617.5 | 0.0 | 38.0 | 30.0 | 967.2 | 793.5 | |
| Kingulungulu | 1 | 123.8 | 223.1 | 0.0 | 6.5 | 0.0 | 353.4 | 288.4 |
| 5 | 261.8 | 468.6 | 11.8 | 7.2 | 0.0 | 749.4 | 609.6 | |
| Kirisirya plantain | 1 | 1.2 | 2.2 | 0.0 | 0.0 | 59.7 | 3.4 | 2.7 |
| 5 | 133.2 | 242.7 | 20.0 | 0.0 | 32.1 | 395.9 | 319.9 | |
| Kothina | 1 | 19.2 | 42.7 | 0.0 | 0.0 | 0.0 | 61.9 | 52.3 |
| 5 | 226.3 | 368.8 | 24.7 | 0.0 | 23.4 | 619.8 | 495.1 | |
| Kothina I | 1 | 13.9 | 29.1 | 0.0 | 0.0 | 0.0 | 43.0 | 36.1 |
| 5 | 204.1 | 453.1 | 22.9 | 0.0 | 9.7 | 680.1 | 567.3 | |
| Kothina II | 1 | 3.4 | 9.5 | 0.0 | 0.0 | 0.0 | 12.9 | 11.2 |
| 5 | 102.9 | 255.6 | 22.7 | 0.0 | 15.3 | 381.2 | 319.1 | |
| Kothina III | 1 | 4.2 | 14.3 | 0.0 | 0.0 | 0.0 | 18.5 | 16.4 |
| 5 | 195.4 | 436.8 | 33.1 | 0.0 | 33.5 | 665.3 | 552.0 | |
| Mabilanga | 1 | 2.4 | 4.7 | 0.0 | 0.0 | 0.0 | 7.1 | 5.4 |
| 5 | 236.6 | 448.2 | 23.0 | 0.0 | 37.2 | 707.8 | 578.7 | |
| Makaka | 1 | 25.5 | 64.8 | 3.2 | 0.0 | 12.1 | 93.5 | 79.2 |
| 5 | 67.1 | 176.8 | 15.4 | 0.0 | 20.3 | 259.3 | 218.5 | |
| Makpelekese | 1 | 6.1 | 13.7 | 0.0 | 0.0 | 0.0 | 19.8 | 16.8 |
| 5 | 141.6 | 307.0 | 18.5 | 0.0 | 41.3 | 467.1 | 387.6 | |
| Mangondi | 1 | 4.5 | 7.8 | 0.0 | 0.0 | 0.0 | 12.3 | 10.0 |
| 5 | 200.3 | 336.7 | 18.8 | 0.0 | 43.6 | 555.8 | 446.8 | |
| Mangondi I | 1 | 4.5 | 7.8 | 0.0 | 0.0 | 0.0 | 12.3 | 10.0 |
| 5 | 200.3 | 336.7 | 18.8 | 0.0 | 43.6 | 555.8 | 446.8 | |
| Mangondi II | 1 | 11.6 | 28.3 | 0.0 | 0.0 | 0.0 | 39.9 | 34.1 |
| 5 | 229.2 | 353.2 | 15.9 | 0.0 | 39.8 | 598.3 | 476.2 | |
| Manzenzele | 1 | 6.9 | 11.0 | 0.0 | 0.0 | 3.7 | 17.9 | 14.5 |
| 5 | 103.1 | 187.7 | 13.3 | 0.0 | 22.8 | 304.1 | 246.3 | |
| Mayayi | 1 | 11.3 | 23.6 | 0.0 | 0.0 | 0.0 | 34.9 | 29.3 |
| 5 | 328.8 | 631.0 | 43.7 | 0.0 | 44.7 | 1003.5 | 818.5 | |
| Musilongo | 1 | 4.1 | 11.1 | 0.0 | 0.0 | 0.0 | 15.2 | 12.4 |
| 5 | 145.6 | 308.2 | 18.3 | 0.0 | 23.9 | 472.1 | 390.7 | |
| Ndonge | 1 | 4.6 | 8.3 | 0.0 | 0.0 | 0.0 | 12.9 | 10.5 |
| 5 | 163.7 | 336.8 | 21.4 | 0.0 | 21.9 | 521.9 | 430.0 | |
| Ngobia II‐1 | 1 | 47.2 | 122.6 | 0.0 | 23.0 | 3.2 | 192.8 | 147.4 |
| 5 | 98.9 | 235.5 | 18.0 | 28.6 | 19.3 | 381.0 | 309.7 | |
| Ngobia mukelekele | 1 | 4.0 | 5.0 | 0.0 | 0.0 | 0.0 | 9.0 | 7.0 |
| 5 | 249.5 | 376.3 | 18.5 | 0.0 | 47.5 | 644.3 | 510.8 | |
| Nguma | 1 | 34.4 | 88.4 | 0.0 | 2.8 | 0.0 | 125.6 | 107.1 |
| 5 | 267.5 | 529.4 | 0.0 | 32.4 | 26.0 | 829.3 | 680.3 | |
| Nguma II | 1 | 26.9 | 59.3 | 2.7 | 0.0 | 16.0 | 88.9 | 74.2 |
| 5 | 164.1 | 344.3 | 22.1 | 0.0 | 40.3 | 530.5 | 438.1 | |
| Nguma III | 1 | 12.6 | 31.4 | 0.0 | 0.0 | 0.0 | 44.0 | 37.7 |
| 5 | 184.0 | 338.9 | 22.1 | 0.0 | 29.3 | 545.0 | 442.6 | |
| Nzirabahima | 1 | 23.4 | 30.3 | 1.0 | 0.0 | 1.9 | 54.7 | 42.5 |
| 5 | 521.6 | 645.3 | 33.2 | 0.0 | 21.5 | 1200.1 | 923.7 | |
| Plantain masunga | 1 | 25.6 | 51.0 | 0.0 | 0.0 | 0.0 | 76.6 | 63.9 |
| 5 | 234.6 | 494.1 | 10.5 | 13.0 | 13.0 | 752.2 | 623.9 | |
| Plantain grand format I | 1 | 71.9 | 2.9 | 0.0 | 0.0 | 0.0 | 74.8 | 38.8 |
| 5 | 94.9 | 165.1 | 10.6 | 0.0 | 28.0 | 270.6 | 218.0 | |
| Plantain grand format II | 1 | 33.1 | 72.9 | 4.9 | 0.0 | 3.6 | 110.9 | 92.1 |
| 5 | 103.0 | 232.0 | 20.8 | 0.0 | 20.2 | 355.8 | 294.5 | |
| Sanza moya | 1 | 10.9 | 37.8 | 0.0 | 0.0 | 0.0 | 48.7 | 28.1 |
| 5 | 113.0 | 331.6 | 8.1 | 23.9 | 3.9 | 476.6 | 405.1 | |
| Sanza tatu | 1 | 24.8 | 34.1 | 0.0 | 0.0 | 0.0 | 58.9 | 46.5 |
| 5 | 363.2 | 552.2 | 13.0 | 0.0 | 10.1 | 928.4 | 740.7 | |
| UCG II | 1 | 183.2 | 351.4 | 13.6 | 0.0 | 18.2 | 548.2 | 450.2 |
| 5 | 91.1 | 162.6 | 8.8 | 0.0 | 17.3 | 262.5 | 212.8 | |
| UCG III | 1 | 8.1 | 9.8 | 0.0 | 0.0 | 0.0 | 17.9 | 213.8 |
| 5 | 253.4 | 367.4 | 29.6 | 0.0 | 42.4 | 650.4 | 509.8 | |
| UCG IV | 1 | 13.6 | 20.8 | 0.0 | 0.0 | 0.0 | 34.4 | 27.6 |
| 5 | 207.4 | 407.3 | 0.0 | 0.0 | 9.4 | 614.7 | 511.0 | |
| UCG VIII | 1 | 11.9 | 26.5 | 0.0 | 0.0 | 0.0 | 38.4 | 32.4 |
| 5 | 99.2 | 213.3 | 19.9 | 0.0 | 20.7 | 332.4 | 273.4 | |
| Vuhembe | 1 | 43.5 | 82.6 | 0.0 | 0.0 | 4.4 | 126.1 | 104.4 |
| 5 | 147.8 | 285.0 | 16.2 | 0.0 | 34.2 | 449.0 | 367.5 | |
| Vuhetera | 1 | 17.3 | 46.5 | 0.0 | 0.0 | 0.0 | 63.8 | 55.1 |
| 5 | 234.1 | 499.8 | 0.0 | 37.0 | 24.3 | 770.9 | 636.4 | |
| Vuhindi | 1 | 2.7 | 4.0 | 0.0 | 0.0 | 0.0 | 6.7 | 5.3 |
| 5 | 91.1 | 163.2 | 12.6 | 0.0 | 17.6 | 266.9 | 215.4 | |
| Vulambya | 1 | 40.9 | 106.7 | 0.0 | 0.0 | 0.0 | 147.6 | 127.2 |
| 5 | 174.0 | 418.3 | 0.0 | 20.3 | 0.0 | 612.6 | 516.1 | |
| LSD (5%) | 64.8 | 112.5 | 7.7 | 6.5 | 27.4 | 187.9 | 147.4 | |
| P value | <0.001 | <0.001 | <0.001 | <0.001 | <0.001 | <0.001 | <0.001 |
t‐AC, all‐trans‐a‐carotene; t‐BC, all‐trans‐b‐carotene; cis‐AC, cis ‐a‐carotene; cis‐BC, cis ‐b‐carotene.
Figure 1.

Proportion (%) of different pro‐vitamin A carotenoids at ripening stage 1 for the assessed plantain landraces.
Figure 2.

Proportion (%) of different pro‐vitamin A carotenoids at ripening stage 5 for the assessed plantain landraces.
A high variability (P < 0.001) in the total pVACs content was observed amongst the 48 plantain cultivars and between the two ripening stages. The mean total pVACs content varied from 4 μg/100 gfw in ‘Akange’ to 548 μg/100 gfw in ‘UCG II’ at ripening stage 1, and from 175 μg/100 gfw in ‘Akobanzi’ to 1756 μg/100 gfw in ‘Adili II’ at stage 5 (Table 3). Wide variations in pVACs content were also reported for Asian cultivars (300 to 6360 μg/100 gfw) within and across banana genomic groups and for African cultivars30, 34, 35, 38 (Table 1).
Significant differences (P < 0.001) were observed between the various cultivars and the two ripening stages (Table 3). Except for ‘UCG II’ in which the total pVACs decreased from 548 to 263 μg/100 gfw with ripening, a (predominantly significant) increase in the t‐BC, t‐AC, total pVACs, and t‐BCE levels from ripening stage 1 to 5 was observed for all the cultivars (Table 3). For example, the largest increments in fruit pulp mean total pVACs of 118, 75, 72, 47, 45 and 45‐fold were observed for ‘Mabilanga’, ‘Akoto Membo’, ‘Ngobia Mukelekele’, ‘Adili II’, ‘Mangondi’ and ‘Mangondi I’, and ‘Akange’, respectively (Table 3). Similar increases in pVACs from ripening stage 1 to 5 in different banana cultivars were observed by Ekesa et al. (2013, 2015)33, 39, 47 and Newilah et al. (2008),63 which are attributed to the enhanced carotenogenesis that commonly occurs during maturation or ripening of fruits and vegetables.12, 61, 63
At ripening stage 5, 29 plantain cultivars (i.e. 60% of the studied cultivars) had a mean total pVACs content higher than 500 μg/100 gfw (i.e. RAE of >32 μg/100 gfw) while three cultivars had a mean total pVACs value above 1000 μg/100 gfw, with the highest value of 1756 μg/100 gfw recorded in the cultivar ‘Adili II’ (Tables 3 and 4). The quantities of pVACs measured in these DR Congolese plantains (ranging from 175 to 1756 μg/100 gfw, with BCE values between 147 and 1359 μg/100 gfw) are higher than those reported for East African highland and dessert banana cultivars,36, 40 while they are comparable to most Pacific (Musa Iholena and Maoli‐Pōpō‘ulu subgroups) and other African plantains34, 35, 36, 37, 38, 39, 46, 64; cf. Tables 1 and 3). These values are, however, significantly lower than, for example, the highest measured pVACs content in carrots (11 427 to 14 693 μg/100 gfw), tomatoes (3454 μg/100 gfw), spinach (9890 μg/100 gfw), orange‐fleshed sweet potato (8329–10 699 μg/100 gfw), the Micronesian Fe'i Musa spp. cultivar ‘Utimwas’ (9000 μg/100 gfw), and the Pacific plantain (AAB) cultivar ‘Mangat en Seipahn’ (8207 μg/100 gfw) (Table 1).
Table 4.
Mean fruit pulp retinol activity equivalent (RAE) and the percentage contribution of the pVACs in 48 plantain (AAB) cultivars at ripening stage 1 and 5 toward the dietary reference intakes (DRIs) for children (1–5 years) and women of reproductive age
| % DRI Child (<5 years) | %DRI woman | ||||||
|---|---|---|---|---|---|---|---|
| Cultivar name | Ripening stage | RAE μg/100gfw | Mean in 100 g of fruit pulp | Mean in 250 g of fruit pulp | Mean in 100 g of fruit pulp | Mean in 250 g of fruit pulp | Mean in 500 g of fruit pulp |
| Adili I | 1 | 20.2 | 5.0 | 12.6 | 2.9 | 7.2 | 14.4 |
| 5 | 38.5 | 9.6 | 24.0 | 5.5 | 13.7 | 27.5 | |
| Adili II | 1 | 2.4 | 0.6 | 1.5 | 0.4 | 0.9 | 1.7 |
| 5 | 113.2 | 28.3 | 70.8 | 16.2 | 40.4 | 80.9 | |
| Agbindolo | 1 | 16.8 | 4.2 | 10.5 | 2.4 | 6.0 | 12.0 |
| 5 | 36.8 | 9.2 | 23.0 | 5.3 | 13.1 | 26.3 | |
| Akange | 1 | 0.3 | 0.1 | 0.2 | 0.1 | 0.1 | 0.2 |
| 5 | 12.9 | 3.2 | 8.1 | 1.8 | 4.6 | 9.2 | |
| Akobanzi | 1 | 0.8 | 0.2 | 0.5 | 0.1 | 0.3 | 0.6 |
| 5 | 12.2 | 3.1 | 7.6 | 1.7 | 4.4 | 8.7 | |
| Akoto membo | 1 | 0.9 | 0.2 | 0.5 | 0.1 | 0.3 | 0.6 |
| 5 | 57.7 | 14.4 | 36.1 | 8.2 | 20.6 | 41.2 | |
| Akoto monama | 1 | 3.6 | 0.9 | 2.2 | 0.5 | 1.3 | 2.6 |
| 5 | 44.5 | 11.1 | 27.8 | 6.4 | 15.9 | 31.8 | |
| Alongo | 1 | 1.7 | 0.4 | 1.0 | 0.2 | 0.6 | 1.2 |
| 5 | 37.3 | 9.3 | 23.3 | 5.3 | 13.3 | 26.6 | |
| Apakumo | 1 | 10.8 | 2.7 | 6.7 | 1.5 | 3.8 | 7.7 |
| 5 | 29.9 | 7.5 | 18.7 | 4.3 | 10.7 | 21.3 | |
| Ayaya | 1 | 2.0 | 0.5 | 1.3 | 0.3 | 0.7 | 1.5 |
| 5 | 29.5 | 7.4 | 18.5 | 4.2 | 10.5 | 21.1 | |
| Bakpulu | 1 | 2.6 | 0.7 | 1.6 | 0.4 | 0.9 | 1.9 |
| 5 | 24.6 | 6.2 | 15.4 | 3.5 | 8.8 | 17.6 | |
| Bubu | 1 | 4.7 | 1.2 | 2.9 | 0.7 | 1.7 | 3.3 |
| 5 | 54.0 | 13.5 | 33.8 | 7.7 | 19.3 | 38.6 | |
| Buembe | 1 | 8.1 | 2.0 | 5.1 | 1.2 | 2.9 | 5.8 |
| 5 | 66.1 | 16.5 | 41.3 | 9.5 | 23.6 | 47.2 | |
| Kingulungulu | 1 | 24.0 | 6.0 | 15.0 | 3.4 | 8.6 | 17.2 |
| 5 | 50.8 | 12.7 | 31.8 | 7.3 | 18.1 | 36.3 | |
| Kirisiriya plantain | 1 | 2.3 | 0.1 | 0.1 | 0.0 | 0.1 | 0.2 |
| 5 | 26.7 | 6.7 | 16.7 | 3.8 | 9.5 | 19.0 | |
| Kothina | 1 | 4.4 | 1.1 | 2.7 | 0.6 | 1.6 | 3.1 |
| 5 | 41.3 | 10.3 | 25.8 | 5.9 | 14.7 | 29.5 | |
| Kothina I | 1 | 3.0 | 0.8 | 1.9 | 0.4 | 1.1 | 2.2 |
| 5 | 47.3 | 11.8 | 29.5 | 6.8 | 16.9 | 33.8 | |
| Kothina II | 1 | 0.9 | 0.2 | 0.6 | 0.1 | 0.3 | 0.7 |
| 5 | 26.6 | 6.7 | 16.6 | 3.8 | 9.5 | 19.0 | |
| Kothina III | 1 | 1.4 | 0.3 | 0.9 | 0.2 | 0.5 | 10.0 |
| 5 | 46.0 | 11.5 | 28.8 | 6.6 | 16.4 | 32.9 | |
| Mabilanga | 1 | 0.5 | 0.1 | 0.3 | 0.1 | 0.2 | 0.3 |
| 5 | 48.2 | 12.1 | 30.1 | 6.9 | 17.2 | 34.5 | |
| Makaka | 1 | 6.6 | 1.7 | 4.1 | 0.9 | 2.4 | 4.7 |
| 5 | 18.2 | 4.6 | 11.4 | 0.3 | 6.5 | 13.0 | |
| Makpelekese | 1 | 1.4 | 0.4 | 0.9 | 0.2 | 0.5 | 1.0 |
| 5 | 32.3 | 8.1 | 20.2 | 4.6 | 11.5 | 23.1 | |
| Mangondi | 1 | 0.8 | 0.2 | 0.5 | 0.1 | 0.3 | 0.6 |
| 5 | 37.2 | 9.3 | 23.3 | 5.3 | 13.3 | 26.6 | |
| Mangondi I | 1 | 0.8 | 0.2 | 0.5 | 0.1 | 0.3 | 0.6 |
| 5 | 37.2 | 9.3 | 23.3 | 5.3 | 13.3 | 26.6 | |
| Mangondi II | 1 | 2.8 | 0.7 | 1.8 | 0.4 | 1.0 | 2.0 |
| 5 | 39.7 | 9.9 | 24.8 | 5.7 | 14.2 | 28.4 | |
| Manzenzele | 1 | 1.2 | 0.3 | 0.8 | 0.2 | 0.4 | 0.9 |
| 5 | 20.5 | 5.1 | 12.8 | 2.9 | 7.3 | 14.7 | |
| Mayayi | 1 | 2.4 | 0.6 | 1.5 | 0.4 | 0.9 | 1.7 |
| 5 | 68.2 | 17.1 | 42.6 | 9.7 | 24.4 | 48.7 | |
| Musilongo | 1 | 1.0 | 0.3 | 0.7 | 0.2 | 0.4 | 0.7 |
| 5 | 32.6 | 8.1 | 20.4 | 4.7 | 11.6 | 23.3 | |
| Ndonge | 1 | 0.9 | 0.2 | 0.6 | 0.1 | 0.3 | 0.6 |
| 5 | 35.8 | 9.0 | 22.4 | 5.1 | 12.8 | 25.6 | |
| Ngobia II‐1 | 1 | 12.3 | 3.1 | 7.7 | 1.8 | 4.4 | 8.8 |
| 5 | 25.8 | 6.5 | 16.1 | 3.7 | 9.2 | 18.4 | |
| Ngobia mukelekele | 1 | 0.6 | 0.2 | 0.4 | 0.1 | 0.2 | 0.4 |
| 5 | 42.6 | 10.6 | 26.6 | 6.1 | 15.2 | 30.4 | |
| Nguma | 1 | 8.9 | 2.2 | 5.6 | 1.3 | 3.2 | 6.4 |
| 5 | 56.7 | 14.2 | 35.4 | 8.1 | 20.2 | 40.5 | |
| Nguma II | 1 | 6.2 | 1.6 | 3.9 | 0.9 | 2.2 | 4.4 |
| 5 | 36.5 | 9.1 | 22.8 | 5.2 | 13.0 | 26.1 | |
| Nguma III | 1 | 3.1 | 0.8 | 2.0 | 0.5 | 1.1 | 2.2 |
| 5 | 36.9 | 9.2 | 23.1 | 5.3 | 13.2 | 26.4 | |
| Nzirabahima | 1 | 3.5 | 0.9 | 2.2 | 0.5 | 1.3 | 2.5 |
| 5 | 77.0 | 19.2 | 48.1 | 11.0 | 27.5 | 55.0 | |
| Plantain masunga | 1 | 5.3 | 1.3 | 3.3 | 0.8 | 1.9 | 3.8 |
| 5 | 52.0 | 13.0 | 32.5 | 7.4 | 18.6 | 37.1 | |
| Plantain grand format I | 1 | 3.2 | 0.8 | 2.0 | 0.5 | 1.2 | 2.3 |
| 5 | 18.2 | 4.5 | 11.4 | 2.6 | 6.5 | 13.0 | |
| Plantain grand format II | 1 | 7.7 | 1.9 | 4.8 | 11.0 | 2.7 | 5.5 |
| 5 | 24.5 | 6.1 | 15.3 | 3.5 | 8.8 | 17.5 | |
| Sanza moya | 1 | 2.3 | 0.6 | 1.5 | 0.3 | 0.8 | 1.7 |
| 5 | 33.8 | 8.4 | 21.1 | 4.8 | 12.1 | 24.1 | |
| Sanza tatu | 1 | 3.9 | 1.0 | 2.4 | 0.5 | 1.4 | 2.8 |
| 5 | 61.7 | 15.4 | 38.6 | 8.8 | 22.0 | 44.1 | |
| UCG II | 1 | 37.5 | 9.4 | 23.5 | 5.4 | 13.4 | 26.8 |
| 5 | 17.7 | 4.4 | 11.1 | 2.5 | 6.3 | 12.7 | |
| UCG III | 1 | 1.2 | 0.3 | 0.7 | 0.2 | 0.4 | 0.8 |
| 5 | 42.5 | 10.6 | 26.6 | 6.1 | 15.2 | 30.4 | |
| UCG IV | 1 | 2.3 | 0.6 | 1.4 | 0.3 | 0.8 | 1.6 |
| 5 | 42.6 | 10.7 | 26.6 | 6.1 | 15.2 | 30.4 | |
| UCG VIII | 1 | 2.7 | 0.7 | 1.7 | 0.4 | 1.0 | 1.9 |
| 5 | 22.8 | 5.7 | 14.2 | 3.3 | 8.1 | 16.3 | |
| Vuhembe | 1 | 8.7 | 2.2 | 5.4 | 1.2 | 3.1 | 6.2 |
| 5 | 30.6 | 7.7 | 19.1 | 4.4 | 10.9 | 21.9 | |
| Vuhetera | 1 | 4.6 | 1.2 | 2.9 | 0.7 | 1.6 | 3.3 |
| 5 | 53.0 | 13.3 | 33.2 | 7.6 | 18.9 | 37.9 | |
| Vuhindi | 1 | 0.4 | 0.1 | 0.3 | 0.1 | 0.2 | 0.3 |
| 5 | 18.0 | 4.5 | 11.2 | 2.6 | 6.4 | 12.8 | |
| Vulambya | 1 | 10.6 | 2.7 | 6.6 | 1.5 | 3.8 | 7.6 |
| 5 | 43.0 | 10.8 | 26.9 | 6.1 | 15.4 | 30.7 | |
| Lsd (5%) | 12.3 | 3.1 | 7.7 | 1.8 | 4.4 | 8.8 | |
| P‐value | <0.001 | <0.001 | <0.001 | <0.001 | <0.001 | <0.001 | |
Values are means of three individual samples per ripening stage per cultivar.
The Dietary Reference Intakes (DRIs) of vitamin A for children 1–5 years old and women of reproductive age is 400 μg RAE and 700 μg RAE, respectively (FAO/WHO, 2002).60
The RAE values of banana cultivars studied at ripening stage 1 and 5 ranged from 0.3 to 37.5 μg/100 gfw and 12 to 113 μg/100 gfw, respectively (Table 4). Englberger et al. (2006)37 reported RAE values of 130 μg/100 gfw for the Fe'i cultivar ‘Asupina’, 79 μg/100 gfw for a horn plantain, and 49 μg/100 gfw for a Pacific plantain. Englberger et al. (2006)64 reported that Musa cultivars with RAE values higher than 44 μg/100 gfw could be considered as excellent sources of pVACS. The RAE values recorded in ‘Adili II’ (113 μg/100 gfw) and ‘Nzirabahima’ (77 μg/100 gfw) exceeded 70 at stage 5, whereas three additional cultivars, ‘Mayayi’, ‘Buembe’ and ‘Sanza Tatu’ had RAE values between 60 and 70 at stage 5, and can thus be considered as highly promising cultivars. An additional ten cultivars had RAE values of 44 μg/100 gfw and above and could also be considered as promising (Table 4).
Daily consumption of 250 g of unprocessed fruit pulp of the cultivars under study at ripening stage 5 meets between 7.6% and 71.8% of the DRI for vitamin A of children below 5 years old (Table 4). Consumption of 500 g of fruit pulp meets between 8.7% and 81% of the DRI for vitamin A of women of reproductive age. The current results show that none of the plantain cultivars can meet 100% of the DRI for vitamin A of the target group when normal amounts of pulp are consumed (Table 4). However, in the east and central African highland region, daily consumption levels of above 500 g have also been reported.27, 29 Slight increments in consumption of some of the plantain cultivars above the normal consumption levels (i.e. 250 g in children below 5 years and 500 g in women of reproductive age) would profoundly increase the percentage of the DRI of vitamin A met. For example, for ‘Adili II’, a daily consumption of 618 g translates to 100% of the DRI for vitamin A in women of reproductive age.
In this study, the pVACs values used to estimate contribution to the DRIs are of raw plantains. Plantains in North Kivu, however, are mainly consumed boiled or fried / roasted.47 It has been established that preparation methods affect the amount and bio‐accessibility of pVACs.32, 43, 65, 66 For example, Ekesa et al. (2013)39, 47 observed total pVACs retention levels of 40–90% and >95% in boiled EAHB and plantains, respectively, and 100% retention when plantains were deep‐fried in fully refined palm oil or crude red palm oil for 2 min. Deep frying with fully refined palm oil and crude red palm oil (with an extremely high pVACs content; see also Table 1) enhanced the BCE and RAE of plantain‐based dishes by twofold and sixfold, respectively.33 Promoting appropriate and healthy fruit processing methods could increase the RAE values of plantain‐based dishes to meet the DRI requirements under normal consumption levels for both children below 5 years and women of reproductive age. Ekesa et al. (2015)33 also observed the bio‐accessibility of pVACs in boiled bananas and plantains to vary with the cultivar and the type of pVACs. These authors noted a significantly more efficient micellarization of t‐BC in the boiled EAHB ‘Vulambya’ (28.9%) than in boiled plantain ‘Musilongo’ (16.6%) and similar bio‐accessibility levels for t‐BC (17–29%) and t‐AC (16–31%). The actual quantity of carotenoids available in these plantains after preparation and consumption therefore still needs to be determined.
Yield of the plantain cultivars and their relative abundance
Significant differences (P < 0.001) were observed in the mean time to harvest (days), bunch weight (kg) and annual yield (t/ha/year) of the plantain cultivars computed over two crop cycles (i.e. the plant crop and first ratoon crop) (Table 5). Mean time to harvest for the plantain cultivars varied between 349 and 688 days. It is important to note that the production / cultivation site (Butembo) is at a high altitude of 1815 m a.s.l. and has a low mean annual temperature (19 °C). Flowering and harvesting of banana (including plantains) is strongly associated with altitude and temperature.67, 68 However, altitude effects on pVACs content have not yet been studied.
Table 5.
The ranking of 48 plantain cultivars for fruit mean total pro‐vitamin A carotenoid content (pVACs) and their respective mean time to harvest, bunch weight and yields
| Cultivar name | Rank for total pVACs | Mean time to harvest (days) | Bunch weight (kg) | Yield (t/ha/y) |
|---|---|---|---|---|
| Adili II | 1 | 500.8ghi | 24.1nopq | 24.5ijklm |
| Nzirabahima | 2 | 396.0b | 22.0ijkl | 30.7rs |
| Mayayi | 3 | 595.0p | 22.3jklm | 22.2fgh |
| Buembe | 4 | 566.2no | 20.1efgh | 19.6de |
| Sanza tatu | 5 | 504.3hij | 16.7b | 17.3cd |
| Akoto membo | 6 | 532.5klm | 26.6s | 26.4mnop |
| Nguma | 7 | 509.9ijkl | 28.3t | 27.4nopq |
| Bubu | 8 | 554.5mn | 25.5qrs | 24.4ijklm |
| Vuhetera | 9 | 501.3ghi | 21.0fghij | 22.2fgh |
| Plantain masunga | 10 | 588.7op | 22.3jklm | 20.0ef |
| Kingulungulu | 11 | — | — | — |
| Mabilanga | 12 | 466.3def | 25.9rs | 29.5qrs |
| Kothina III | 13 | 465.7def | 21.9ijkl | 23.1ghijk |
| UCG III | 14 | 432.3c | 21.4ghijk | 26.5mnop |
| Ngobia mukelekele | 15 | 541.8mn | 19.9efg | 20.2ef |
| Kothina I | 16 | 478.5efgh | 21.9ijkl | 23.3ghijk |
| Akoto monama | 17 | 528.2ijklm | 12.8a | 13.5ab |
| Adili I | 18 | 505.7hijk | 22.5jklmn | 22.2fgh |
| Kothina | 19 | 451.0cde | 18.2bcd | 21.0efg |
| Mangondi II | 20 | — | — | — |
| UCG IV | 21 | 349.3a | 22.8klmn | 35.4t |
| Vulambya | 22 | 387.2b | 20.0efg | 29.6qrs |
| Mangondi | 23 | 533.1klm | 24.8opqr | 24.2ijklm |
| Mangondi I | 24 | — | — | — |
| Nguma III | 25 | 539.2mn | 26.5s | 25.0jklm |
| Nguma II | 26 | 474.8efg | 26.5s | 28.4pqr |
| Agbindolo | 27 | 600.5p | 17.4bc | 16.2c |
| Alongo | 28 | 593.9op | 23.8mnop | 21.2efgh |
| Ndonge | 29 | 548.8mn | 18.8cde | 16.5c |
| Makpelekese | 30 | 593.5op | 20.9fghij | 20.0ef |
| Musilongo | 31 | 478.2efgh | 24.1nopq | 27.9opq |
| Vuhembe | 32 | 397.4b | 23.2lmno | 31.2s |
| Sanza moya | 33 | 549.8mn | 13.0a | 11.8a |
| Ayaya | 34 | 435.8c | 21.2ghijk | 25.1klmn |
| Kirisirya plantain | 35 | — | — | — |
| Apakumo | 36 | — | — | — |
| Kothina II | 37 | 472.2ef | 19.4def | 21.1efg |
| Ngobia II‐1 | 38 | — | — | — |
| Plantain grand format II | 39 | — | — | — |
| Bakpulu | 40 | — | — | — |
| UCG VIII | 41 | — | — | — |
| Manzenzele | 42 | 531.1jklm | 21.7hijkl | 23.5hijkl |
| Plantain grand format I | 43 | 643.1q | 32.7u | 25.8lmno |
| Vuhindi | 44 | 440.3cd | 20.0efg | 22.7ghij |
| UCG II | 45 | 541.5mn | 25.1pqrs | 27.9opq |
| Makaka | 46 | 530.4klm | 16.4b | 16.2c |
| Akange | 47 | 687.7r | 17.2bc | 14.2bc |
| Akobanzi | 48 | 534.8lm | 20.6fghi | 22.3fghi |
| Lsd (5%) | 27.9 | 1.6 | 2.3 | |
| P value | <0.001 | <0.001 | <0.001 | |
| Cv % | 6.0 | 8.3 | 12.0 |
‘***’ denote a significant difference between the cultivars within a yield attribute at P < 0.001.
Means in the column followed by the same letter are not significantly different at 5% least significant difference (Lsd).
Dash (‘—’) denotes that yield data was not collected for the particular cultivar.
Englberger et al. (2006)37 reported the Fe'i cultivar ‘Utin lap’, which has the highest pVACs content measured to date, grows very slowly, and as such, a high pVACs content has been postulated to be associated with slow plant growth. Similarly, Swennen and Wilson (1983)69 reported that most plantains characterized by orange fruit pulp, and thus likely higher pVACs content, have long intervals between crop cycles. The long cycle duration is attributed to a shortage of gibberellins, which is associated with a high apical dominance and corresponding slow sucker growth. However, in this study, regression analysis did not find a strong association between mean plant growth cycle duration and pVACs content (R2 = 0.05).
Bunch weights varied between 12.8 kg in ‘Akoto monama’ and 33 kg in ‘Plantain Grand Format I’; 82% of the cultivars had acceptable bunch weights of 20 kg and above (Table 5). However, no strong association (R2 = 0.05) was observed between the bunch weights and the total pVACs content across all the cultivars. The mean annual yield of the 48 plantain cultivars varied between 12 t/ha/year in ‘Sanza Moya’ and 35 t/ha/year in ‘UCG IV’, with 82% having yields above 20 t/ha/year (Table 5). The best three cultivars in terms of pVACs concentration, namely ‘Adili II’, ‘Nzirabahima’ and ‘Mayayi’, recorded annual yields of 25, 31, and 22 t/ha/year, respectively. ‘Nzirabahima’, the second best cultivar in terms of pVACs content, ranked the third in terms of bunch yield. ‘Nguma I’ and ‘Akoto Nembo’, which had moderately high pVACs content, also rated among the cultivars with the best yields. In contrast, the top two performers in terms of bunch yield, ‘UCG IV’ and ‘Vuhembe’, respectively, ranked only 21st and 32nd in terms of pVACs (Table 5).
It is important to note that the three cultivars with the highest pVACs content are not widely distributed or grown in the study region (i.e. eastern DR Congo) (Table 2). This could partially explain the fact that this region still experiences high levels of VAD, especially among children below 5 years and women of reproductive age, despite the presence of such plantain cultivars. There is a need to understand the reasons for the low adoption or growth of these cultivars, and to promote their cultivation and consumption within households deliberately. The most important selection criteria influencing farmers' choices of plantain cultivars to adopt and or retain on farm in the central African region (Burundi, eastern DR Congo and Rwanda) include the taste and flavor of the pulp, size of the bunch, crop‐cycle duration, and market demand.48, 70 Plantain cultivars with small bunch sizes have a lower market value and are more likely to be phased out / rejected by farmers irrespective of the nutritional value, whereas cultivars with average to large bunches are favored by farmers for their high marketability. The observed bunch weights and annual yields, especially for the cultivars with the highest levels of pVACs, are moderately high and would thus positively contribute to scaling‐out efforts. For example, ‘Nzirabahima’ and ‘Nguma’, which respectively ranked second and seventh in vitamin A content, are expected to be highly marketable due to their large bunch sizes and taste. The orientation towards taking such cultivars with large bunches to markets could, however, potentially be an undoing in the fight against VAD, for the poor rural households. Sensitization will also be necessary to change such attitudes and to enhance household consumption of vitamin A‐rich plantains.
CONCLUSION
In this study, 15 plantain cultivars, including ‘Adili II’, ‘Nzirabahima’, ‘Mayayi’, ‘Buembe’, and ‘Sanza Tatu’ (associated with RAE values above 44 μg/100 gfw) can be considered as good sources of pVACs. Modest consumption (250 or 500 gfw) of the fruit pulp of the five best plantain cultivars at ripening stage 5 meets between 39–71% and 44–81% of vitamin A DRI respectively, for children below 5 years old and women of reproductive age. The 15 best plantain cultivars (especially the top five) could potentially be introduced / promoted as alternative sources of pro‐vitamin A in banana‐dependent communities. Given the cultural importance of plantains in diets in the region studied, targeting them in the fight against VAD offers an additional sustainable alternative to complement existing pVACs‐rich crops such as carrots and sweet potato. It is important to note that the best plantain cultivars for pVACs content are currently not widely cultivated and thus need to be actively promoted. Despite the fact that the amount of pVACs in fruit pulp did not have a strong association with crop‐cycle duration and yield (in tons/ha/year), most of the plantain cultivars including the best five in term of pVACs content had moderate to large bunch sizes and yields (tons/ha/year). Bunch size and yield are critical criteria used by farmers in selecting Musa cultivars to grow and retain on farms, and could thus be helpful in promoting the scaling out of these cultivars. Additional studies are needed to further understand the factors responsible for the current often limited geographical spread and potential issues / drivers linked to their introduction / promotion, e.g. pest and disease resistance and culinary / organoleptic characteristics.
ACKNOWLEDGEMENTS
We acknowledge the support of the Directorate General for Development (DGD‐Belgium) for its financial contribution to this work. The authors are grateful to Genáhl Dóra for carrying out the lab analysis at the Catholic University of Leuven (KULeuven). This research was conducted in the framework of the CGIAR Research Program on Roots, Tubers and Bananas (CRP‐RTB).
REFERENCES
- 1. Dowling JE and Wald G, Vitamin A deficiency and night blindness. Proc Natl Acad Sci U S A 44:648–661 (1958). [DOI] [PMC free article] [PubMed] [Google Scholar]
- 2. Sommer A, The journal of nutrition history of nutrition vitamin A deficiency and clinical disease: an historical overview. J Nutr 138:1835–1839 (2008). [DOI] [PubMed] [Google Scholar]
- 3. Zile MH, Function of vitamin A in vertebrate embryonic development. J Nutr 131:705–708 (2001). [DOI] [PubMed] [Google Scholar]
- 4. Maziya‐Dixon BB, Akinyele IO, Sanusi RA, Oguntona TE, Nokoe SK and Harris EW, Vitamin A deficiency is prevalent in children less than 5 y of age in Nigeria. J Nutr 136:2255–2261 (2006). [DOI] [PubMed] [Google Scholar]
- 5. World Health Organization (WHO) , Global Prevalence of Vitamin A Deficiency in Populations at Risk 1995–2005: WHO Global Database on Vitamin A Deficiency. (2009). Available: http://whqlibdoc.who.int/publications/2009/9789241598019_eng.pdf [15 January 2018].
- 6. Ahmed T, Mahfuz M, Ireen S, Ahmed AMS, Rahman S, Islam MM et al, Nutrition of children and women in Bangladesh: trends and directions for the future. J Health Popul Nutr 30:1–11 (2012). [DOI] [PMC free article] [PubMed] [Google Scholar]
- 7. Strobel M, Tinz J and Biesalski HK, The importance of β‐carotene as a source of vitamin A with special regard to pregnant and breastfeeding women. Eur J Nutr 46:1–20 (2007). [DOI] [PubMed] [Google Scholar]
- 8. Humphrey JH, West KP Jr and Sommer A, Vitamin A deficiency and attributable mortality among under‐5‐year‐olds. Bull World Health Organ 70:225–232 (1992). [PMC free article] [PubMed] [Google Scholar]
- 9. SCN, United Nations Standing Committte on Nutrition , Sixth Report on the World Nutrition Situation: Progress in Nutrition. United Nations Standing Committee on Nutrition (SCN), Geneva: (2010). [Google Scholar]
- 10. Van den Berg H, Faulks R, Granado HF, Hirschberg J, Olmedilla B, Sandmann G et al, The potential for the improvement of carotenoid levels in foods and likely systemic effects. J Sci Food Agric 80:880–912 (2000). [Google Scholar]
- 11. Fitzpatrick TB, Basset GJ, Borel P, Carrari F, DellaPenna D, Fraser PD et al, Vitamin deficiencies in humans: can plant science help? Plant Cell 24:395–414 (2012). [DOI] [PMC free article] [PubMed] [Google Scholar]
- 12. Crupi P, Preedy VR and Antonacci D, Cap. 17. HPLC‐DAD‐MS (ESI+) determination of carotenoids in fruit, in Vitamin A: Chemistry, Analysis, Function and Effects, ed. by Preedy V, RSC Publishing, pp. 282–302 (2012). [Google Scholar]
- 13. Devadas RP, Soroja S and Murthy NK, Biological availability of beta carotene from fresh and dried green leafy vegetables on preschool children. Indian J Nutr Diet 15:335 (1978). [Google Scholar]
- 14. Institute of Medicine , Dietary Reference Intakes Report for Vitamin A, Vitamin K, Arsenic, Boron, Chromium, Copper, Iodine, Iron, Manganese, Molybdenum, Nickel, Silicon, Vanadium, and Zinc. Washington, D.C., National Academy Press. (2001). Available: https://www.ncbi.nlm.nih.gov/books/NBK222310/ [15 January 2018].
- 15. Underwood B, Dietary approaches to the control of vitamin A deficiency: an introduction and overview. Food Nutr Bull 21:117–123 (2000). [Google Scholar]
- 16. Tang G, Using plant foods rich in β‐carotene to combat vitamin A deficiency. 13 Sight and Life pp. 20–29. (2013). [Google Scholar]
- 17. Castenmiller JJM and West CE, Bioavailability and bioconversion of carotenoids. Annu Rev Nutr 18:19–38 (1998). [DOI] [PubMed] [Google Scholar]
- 18. Reboul E, Richelle M, Perrot E, Desmoulins‐Malezet C, Pirisi V and Borel P, Bio‐accessibility of carotenoids and vitamin E from their main dietary sources. J Agric Food Chem 54:8749–8755 (2006). [DOI] [PubMed] [Google Scholar]
- 19. Chakravarty I, Food‐based strategies to control vitamin A deficiency. Food Nutr Bull 21:135–143 (2000). [Google Scholar]
- 20. Ministère du Plan et Macro International , Enquête Démographique et de Santé, République Démocratique du Congo 2007. Ministère du Plan et Macro International, Calverton, Maryland: (2008). [Google Scholar]
- 21. Semba RD, Ndugwa C, Perry RT, Clark TD, Jackson JB, Melikian G et al, Effect of periodic vitamin A supplementation on mortality and morbidity of human immunodeficiency virus‐infected children in Uganda: a controlled clinical trial. Nutrition 21:25–31 (2005). [DOI] [PubMed] [Google Scholar]
- 22. Summit , Effect of maternal multiple micronutrient supplementation on fetal loss and infant death in Indonesia: a double‐blind cluster‐randomized trial. The Lancet 371:215–227 (2008). [DOI] [PubMed] [Google Scholar]
- 23. West CE, Meeting requirements for vitamin A. Nutr Rev 58:341–345 (2000). [DOI] [PubMed] [Google Scholar]
- 24. West KP, Vitamin A deficiency disorders in children and women. Food Nutr Bull 24:78–90 (2003). [DOI] [PubMed] [Google Scholar]
- 25. Ekesa BN, Blomme G and Garming H, Dietary diversity and nutritional status of pre‐school children from Musa‐dependent households in Gitega (Burundi) and Butembo (Democratic Republic of Congo). Afr J Food Agric Nutr Dev 11:4896–4911 (2011). [Google Scholar]
- 26. Picq C, Fouré E and Frison EA, Banana and food security. Les productions bananières: un enjeu économique majeur pour la sécurité alimentaire, in Rapport International Symposium. Douala, Cameroun, INIBAP: (1998). [Google Scholar]
- 27. Karamura E, Frison E, Karamura DA and Sharrock S, Banana Production Systems in Eastern and Southern Africa. Bananas and food security. INIBAP, Montpellier, pp. 401–412 (1998). [Google Scholar]
- 28. DRC, Ministry of Planning , Piloting Unit of DSRP Process, Kinshasa‐Gombe. Democratic Republic of Congo, pp. 129–155 (2005). [Google Scholar]
- 29. Edmeades S, Smale M and Karamura D, Biodiversity of bananas on farms in Uganda. Brief 24. Research at a glance. Genetic Resource Policies. Promising Crop Biotechnologies for Smallholder Farmers in East Africa: Bananas and Maize. (2015). Available: https://www.bioversityinternational.org/fileadmin/_migrated/uploads/tx_news/Biodiversity_of_bananas_on_farms_in_Uganda_1207.pdf. [13 January, 2018]
- 30. Davey MW, Garming H, Ekesa B, Roux N and Van den Bergh I, Exploiting banana biodiversity to reduce vitamin A deficiency‐related illness: a fast and cost‐effective strategy, in Human Nutrition and Health Conference, Queensland Primary Industries and Fisheries, Queensland, Australia. 2008, p. 159–171 (2009a). [Google Scholar]
- 31. Davey MW, Saeys W, Hof E, Ramon H, Swennen RL and Keulemans J, Application of visible and near‐infrared reflectance spectroscopy (VIS/NIRS) to determine carotenoids contents in banana (Musa spp.) fruit pulp. J Agric Food Chem 57:1742–1751 (2009b). [DOI] [PubMed] [Google Scholar]
- 32. Ekesa BN, Poulaert M, Davey MW, Kimiywe J, Van den Bergh I, Blomme G et al, Bioaccessibility of provitamin A carotenoids in bananas (Musa spp.) and derived dishes in African countries. Food Chem 133:1471–1477 (2012). [Google Scholar]
- 33. Ekesa B, Nabuuma D, Blomme G and Van den Bergh I, Provitamin A carotenoids content of unripe and ripe banana cultivars for potential adoption in eastern Africa. J Food Compos Anal 43:1–6 (2015). [Google Scholar]
- 34. Englberger L, Aalbersberg W, Ravi P, Bonnin E, Marks GC, Fitzgerald MH et al, Further analyses on Micronesian banana, taro, breadfruit and other foods for provitamin A carotenoids and minerals. J Food Compos Anal. 16:219–236 (2003a). [Google Scholar]
- 35. Englberger L, Darnton‐Hill I, Coyne T, Fitzgerald MH and Marks GC, Carotenoid‐rich bananas: a potential food source for alleviating vitamin A deficiency. Food Nutr Bull 24:303–318 (2003b). [DOI] [PubMed] [Google Scholar]
- 36. Mbabazi R, Molecular Characterization and Carotenoid Quantification of pro‐vitamin A Biofortified Genetically Modified Bananas in Uganda. (2015). Available: http://eprints.qut.edu.au/84853/1/Ruth_Mbabazi_Thesis.pdf.[ 14 January, 2018].
- 37. Englberger L, Shierle J, Aalbersberg W, Hoffmnn P, Humphries J, Huang A et al, Carotenoid and vitamin content of karat and other and other Micronesian banana cultivars. Int J Food Sci Nutr 57:399–418 (2006d). [DOI] [PubMed] [Google Scholar]
- 38. Davey MW, Van den Bergh I, Markham R, Swennen R and Keulemans J, Genetic variability in Musa fruit provitamin A carotenoids, lutein and mineral micronutrient contents. Food Chem 115:806–813 (2009c). [Google Scholar]
- 39. Ekesa BN, Kimiywe J, Van den Bergh I, Blomme G, Dhuique‐Mayer C and Davey M, Content and retention of provitamin A carotenoids following ripening and local processing of four popular Musa cultivars from eastern Democratic Republic of Congo. Sustainable Agric Res 2:60 (2013b). [Google Scholar]
- 40. Fungo R, Kikafunda JK and Pillay M, ß‐carotene, iron and zinc content in Papua New Guinea and east African highland bananas. Afr. J. Food Agric. Nutr. Dev. 10:2629–2644 (2010). [Google Scholar]
- 41. Englberger L, Aalbersberg W, Dolodolotawake U, Schierle J, Humphries J, Iuta T et al, Carotenoids content of pandanus fruit cultivars and other foods of the Republic of Kiribati. Public Health Nutr 9:631–643 (2006a). [DOI] [PubMed] [Google Scholar]
- 42. Englberger L, Aalbersberg W, Schierle J, Marks GC, Fitzgerald MH, Muller F et al, Carotenoids content of different edible pandanus fruit cultivars of the Republic of the Marshall Islands. J Food Compos Anal 19:484–494 (2006b). [Google Scholar]
- 43. van Jaarsveld PJ, De Wet M, Harmse E, Nestel P and Rodriguez‐Amaya DB, Retention of B‐carotene in boiled, mashed orange‐fleshed sweet potato (in: after processing: the fate of food components). J Food Compos Anal 19:321–329 (2006). 10.1016/j.jfca.2004.10.007. [DOI] [Google Scholar]
- 44. Welsch R, Arango J, Bär C, Salazar B, Al‐Babili S, Beltrán J et al, Provitamin A accumulation in cassava (Manihot esculenta) roots driven by a single nucleotide polymorphism in a phytoene synthase gene. Plant Cell 10:3348–3356 (2010). [DOI] [PMC free article] [PubMed] [Google Scholar]
- 45. Tan B, Palm carotenoids, tocopherols and tocotrienols. J Am Oil Chem Soc 66:770–776 (1989). [Google Scholar]
- 46. Davey MW, Stals E, NgohNewilah G, Tomekpe K, Lusty C, Markham R et al, Sampling strategies and variability in fruit pulp micronutrient contents of west and central African bananas and plantains (Musa spp.). J Agric Food Chem 55:2633–2644 (2007). [DOI] [PubMed] [Google Scholar]
- 47. Ekesa BN, Kimiywe J, Davey M, Dhuique‐Mayer C, Van den Bergh I and Blomme G, Contribution of bananas and plantains to the diet and nutrition of Musa‐dependent households with preschoolers in Beni and Bukavu territories, eastern Democratic Republic of Congo, in Banana Systems in the Humid Highlands of Sub‐Saharan Africa Enhancing Resilience and Productivity, ed. by Blomme G, Piet Van Asten P. and Vanlauwe B. CAB International, Wallingford, Oxfordshire, UK. pp. 202–209 (2013a). [Google Scholar]
- 48. Ocimati W, Karamura D, Sivirihauma C, Ndungo V, De Langhe E, Adheka J et al, On‐farm banana (Musa) cultivar diversity status across different altitudes in north and south Kivu provinces of eastern Democratic Republic of Congo, Acta Hort 1114: 35–44 (2016). [Google Scholar]
- 49. Farrow A, Busingye L and Bagenze P, Characterisation of Mandate Areas for the Consortium for Improved Agricultural Livelihoods in Central Africa (CIALCA). CIALCA, Nairobip. 132 (2006). [Google Scholar]
- 50. Ndungo V, Analyse de la sécurité alimentaire menacée en période de guerre dans les territoires de Beni et de Lubero. Parcours et initiatives 1:89–95 (2002). [Google Scholar]
- 51. Sivirihauma C, Ocimati W, Valimuzigha K, Karamura D, Adheka J, Ibanda B et al, Diversity and morphological characterization of Musa spp. in north Kivu and Ituri provinces, eastern Democratic Republic of Congo. Int J Biodiversity Conserv 9:292–305 (2017). [Google Scholar]
- 52. Gaidashova SV, Karemera F and Karamura EB, Agronomic performance of introduced banana varieties in lowlands of Rwanda. Afr Crop Sci J 16:9–16 (2008). [Google Scholar]
- 53. Dadzie BK and Orchard JE, Routine post‐harvest screening of Banana/plantain hybrids: criteria and methods, in INIBAP Technical Guidelines 2. International Plant Genetic Resources Institute, Rome, Italy; International Network for the Improvement of Banana and Plantain, Montpellier, France; ACPEU Technical Centre for Agricultural and Rural Cooperation. Wageningen, The Netherlands, Published by INIBAP. p. 63 (1997). [Google Scholar]
- 54. Stover RH and Simmonds NW, Bananas, in Tropical Agricultural Series, 3rd edn. Longman, Essex, UK: (1987). [Google Scholar]
- 55. Davey MW, Keulemans J and Swennen R, Methods of the efficient quantification of fruit provitamin A contents. J Chromatogr 1136:176–184 (2006). 10.1016/j.chroma.2006.09.077. [DOI] [PubMed] [Google Scholar]
- 56. Shillis ME, Shike M, Ross AC, Caballero B and Cousins RJ, Modern Nutrition in Health and Disease, 10th edn. Lippincott Williams and Wilkins, London, UK (2006). [Google Scholar]
- 57. Trumbo PR, Yates AA, Schlicker‐Renfro S and Suitor C, Dietary reference intakes: revised nutritional equivalents for folate, vitamin E and provitamin A carotenoids. J Food Compos Anal. 16:379–382 (2003). [Google Scholar]
- 58. Fraser PD and Bramley PM, The biosynthesis and nutritional uses of carotenoids. Prog Lipid Res 43:228–265 (2004). [DOI] [PubMed] [Google Scholar]
- 59. Yeum KJ and Russell RM, Carotenoid bioavailability and bioconversion. Annu Rev Nutr 22:483–504 (2002). [DOI] [PubMed] [Google Scholar]
- 60. FAO/WHO , Human Vitamin and Mineral Requirements. Report of a Joint FAO/WHO Expert Consultation, Bangkok, Thailand. Rome, FAO and WHO: (2002). [Google Scholar]
- 61. Rodriguez‐Amaya DB, A Guide to Carotenoid Analysis in Foods. Washington, DC: ILSI Press, 1997. SCN. Sixth Report on the World Nutrition Situation. Progress in Nutrition. United Nations System Standing Committee on Nutrition United Nations System. Geneva, Switzerland: (1997). [Google Scholar]
- 62. VSN International Ltd , GenStat 12th Edition. (2009). Available: http://www.vsni.co.uk.
- 63.Newilah GN, Lusty C, Van den Bergh I, Akyeampong E, Davey M, Tomekpe K. Evaluating bananas and plantains grown in Cameroon as a potential sources of carotenoids. Food 2(2):135–138 (2008). [Google Scholar]
- 64. Englberger L, Wills RB, Blades B, Dufficy L, Daniells JW and Coyne T, Carotenoids content and flesh color of selected banana cultivars growing in Australia. Food Nutr Bull 27:281–291 (2006c). [DOI] [PubMed] [Google Scholar]
- 65. Masrizal MA, Giraud DW and Driskell JA, Retention of vitamin C, iron and β‐carotene in vegetables prepared using different cooking methods. J Food Qual 20:403–418 (2007). [Google Scholar]
- 66. PinheiroSant' Ana HM, Stringheta PC, Brandao SCC and Cordeiro de Azeredo RM, Carotenoid retention and vitamin A value in carrot (Daucus carota L.) prepared by food service. Food Chem 61:145–151 (1998). [Google Scholar]
- 67. Sikyolo I, Sivirihauma C, Ndungo V, De Langhe E, Ocimati W and Blomme G, Growth and yield of plantain cultivars at four sites of differing altitude in north Kivu, eastern Democratic Republic of Congo, in Banana Systems in the Humid Highlands of Sub‐Saharan Africa, CABI, Wallingford, Oxfordshire, UK. p. 48 (2013). [Google Scholar]
- 68. Turner DW, Fortescue JA, Ocimati W and Blomme G, Plantain cultivars (Musa spp. AAB) grown at different altitudes demonstrate cool temperature and photoperiod responses relevant to genetic improvement. Field Crop Res 194:103–111 (2016). [Google Scholar]
- 69. Swennen R and Wilson GF, The stimulation of sword sucker development by applied gibberellin (Ga3) in plantain (Musa spAAB). Fruits 38:261–265 (1983). [Google Scholar]
- 70. Ocimati W, Blomme G, Rutikanga A, Karamura D, Ragama P, Gaidashova S et al, Musa germplasm diversity status across a wide range of agro‐ecological zones in Rwanda. J Appl Biosci 73:5979–5990 (2014). [Google Scholar]
