Cassava Nutrition and Biofortification
Cassava Nutrition and Biofortification
of Cassava for
                                                                               Use as a Staple
                                                                              Food and Recent
                                                                                 Advances for
                                                                                 Improvement
                                                                                                   Julie A. Montagnac, Christopher R. Davis,
                                                                                                                and Sherry A. Tanumihardjo
     ABSTRACT: Cassava is a drought-tolerant, staple food crop grown in tropical and subtropical areas where many
     people are afflicted with undernutrition, making it a potentially valuable food source for developing countries. Cas-
     sava roots are a good source of energy while the leaves provide protein, vitamins, and minerals. However, cassava
     roots and leaves are deficient in sulfur-containing amino acids (methionine and cysteine) and some nutrients are
     not optimally distributed within the plant. Cassava also contains antinutrients that can have either positive or ad-
     verse effects on health depending upon the amount ingested. Although some of these compounds act as antioxidants
     and anticarcinogens, they can interfere with nutrient absorption and utilization and may have toxic side effects. Ef-
     forts to add nutritional value to cassava (biofortification) by increasing the contents of protein, minerals, starch,
     and -carotene are underway. The transfer of a 284 bp synthetic gene coding for a storage protein rich in essential
     amino acids and the crossbreeding of wild-type cassava varieties with Manihot dichotoma or Manihot oligantha
     have shown promising results regarding cassava protein content. Enhancing ADP glucose pyrophosphorylase ac-
     tivity in cassava roots or adding amylase to cassava gruels increases cassava energy density. Moreover, carotenoid-
     rich yellow and orange cassava may be a foodstuff for delivering provitamin A to vitamin A–depleted populations.
     Researchers are currently investigating the effects of cassava processing techniques on carotenoid stability and iso-
     merization, as well as the vitamin A value of different varieties of cassava. Biofortified cassava could alleviate some
     aspects of food insecurity in developing countries if widely adopted.
C   2009 Institute of Food Technologists
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                                                  Vol. 8, 2009—COMPREHENSIVE REVIEWS IN FOOD SCIENCE AND FOOD SAFETY                           181
CRFSFS: Comprehensive Reviews in Food Science and Food Safety
Table 1 --- Maximum recorded yield and food energy of basis (Buitrago 1990) and between 0.4 and 1.5 g/100 g FW (Brad-
important tropical staple crops.a                     bury and Holloway 1988). In contrast, maize and sorghum have
                                                                                about 10 g protein/100 g FW. The content of some essential amino
                                      Annual yield    Daily energy production
                                                                                acids, such as methionine, cysteine, and tryptophan, is very low
Crop                                 (tons/hectare)         (kJ/hectare)
                                                                                (Table 4). However, the roots contain an abundance of arginine,
Fresh cassava root                        71                   1045             glutamic acid, and aspartic acid (Gil and Buitrago 2002). About
Maize grainb                              20                    836             50% of the crude protein in the roots consists of whole protein
Fresh sweet potato root                   65                    752             and the other 50% is free amino acids (predominantly glutamic
Rice grain                                26                    652             and aspartic acids) and nonprotein components such as nitrite,
Sorghum grain                             13                    477             nitrate, and cyanogenic compounds. The presence of cyanogenic
Wheat grain                               12                    460             compounds, which predominate in bitter varieties, and processes
Banana fruit                              39                    334             to reduce them were recently reviewed by Montagnac and others
a Adapted from EL-Sharkawy
                                                                                (2009).
                                (2003).
b All grains reported as dry.
                                                                                Minerals and vitamins
                                                                                    Cassava roots have calcium, iron, potassium, magnesium, cop-
Micronutrients present in staple crops that are being targeted for              per, zinc, and manganese contents comparable to those of many
biofortification include vitamin A, iron, and zinc. Cassava has                 legumes, with the exception of soybeans (Table 5). The calcium
been targeted for biofortification because of its unique geograph-              content is relatively high compared to that of other staple crops
ical distribution and its importance as a staple food. This review              and ranges between 15 and 35 mg/100 g edible portion. The
describes the nutritional value and improvements that researchers               vitamin C (ascorbic acid) content is also high and between 15 to
have achieved in the cassava plant.                                             45 mg/100 g edible portions (Okigbo 1980; Charles and others
                                                                                2004). Cassava roots contain low amounts of the B vitamins, that
                                                                                is, thiamin, riboflavin, and niacin (Table 6), and part of these nu-
Nutritional Value of Cassava Roots                                              trients is lost during processing. Usually the mineral and vitamin
    The composition of cassava depends on the specific tissue (root             contents are lower in cassava roots than in sorghum and maize
or leaf) and on several factors, such as geographic location, va-               (Gil and Buitrago 2002).
riety, age of the plant, and environmental conditions. The roots                    The protein, fat, fiber, and minerals are found in larger quan-
and leaves, which constitute 50% and 6% of the mature cassava                   tities in the root peel than in the peeled root. However, the car-
plant, respectively, are the nutritionally valuable parts of cassava            bohydrates, determined by the nitrogen-free extract, are more
(Tewe and Lutaladio 2004). The nutritional value of cassava roots               concentrated in the peeled root (central cylinder or pulp) (Gil and
is important because they are the main part of the plant consumed               Buitrago 2002). Thus, cassava roots are rich in calories but low
in developing countries. In Table 2, the proximate, mineral, and                in protein, fat, and some minerals and vitamins. Their nutritional
vitamin compositions of cassava roots and leaves are reported. In               value is, consequently, lower than those of cereals, legumes, and
Table 3, the nutrient composition of raw cassava is compared to                 some other root and tuber crops.
other staple crops, such as wheat and corn, and some vegetable
and animal foods.                                                    Processing effects on nutritional value
                                                                       Processing cassava can affect the nutritional value of cassava
Macronutrients                                                       roots through modification and losses in nutrients of high value.
    Cassava root is an energy-dense food. In this regard, cassava Traditional processing techniques and several edible forms of
shows very efficient carbohydrate production per hectare. It pro- cassava roots are illustrated in Figure 2. Analysis of the nutrient
duces about 250000 calories/hectare/d, which ranks it before retention for each cassava edible product (Table 7) shows that raw
maize, rice, sorghum, and wheat (Okigbo 1980). The root is and boiled cassava root keep the majority of high-value nutrients
a physiological energy reserve with high carbohydrate content, except riboflavin and iron. Gari is a common root product that
which ranges from 32% to 35% on a fresh weight (FW) basis, involves grating, fermenting, and roasting. Gari and products ob-
and from 80% to 90% on a dry matter (DM) basis. Eighty percent tained after retting of cassava root with peel are less efficient than
of the carbohydrates produced is starch (Gil and Buitrago 2002); boiled root in keeping nutrients of high value but are better than
83% is in the form of amylopectin and 17% is amylose (Rawel products obtained after retting of shucked cassava roots. How-
and Kroll 2003). Roots contain small quantities of sucrose, glu- ever, the latter is richer in riboflavin than sun-dried flour. Fufu, an
cose, fructose, and maltose (Tewe and Lutaladio 2004). Cassava important staple in Africa, is a mashed cassava root product that
has bitter and sweet varieties. In sweet cassava varieties, up to is allowed to ferment with Lactobacillus bacteria (Sanni and oth-
17% of the root is sucrose with small amounts of dextrose and ers 2002). Medua-me-mbong is a root product that requires only
fructose (Okigbo 1980; Charles and others 2005). Raw cassava boiling and prolonged washing. However, medua-me-mbong has
root has more carbohydrate than potatoes and less carbohydrate the poorest nutritional value compared to other cassava products
than wheat, rice, yellow corn, and sorghum on a 100-g basis with the exception of calcium content (Favier 1977).
(Table 3). The fiber content in cassava roots depends on the vari-     In contrast to boiled cassava, processed root loses a major part
ety and the age of the root. Usually its content does not exceed of dry matter, carbohydrates, protein, and thus calories. Although
1.5% in fresh root and 4% in root flour (Gil and Buitrago 2002). raw cassava root contains significant vitamin C, it is very sensitive
    The lipid content in cassava roots ranges from 0.1% to 0.3% on to heat and easily leaches into water, and therefore almost all of
a FW basis. This content is relatively low compared to maize and the processing techniques seriously affect its content. Boiled cas-
sorghum, but higher than potato and comparable to rice (Table 3). sava, gari, and products resulting from retting of cassava root with
The lipids are either nonpolar (45%) or contain different types of peel, retain thiamin and niacin better than products obtained after
glycolipids (52%) (Hudson and Ogunsua 1974). The glycolipids retting of shucked cassava roots, smoked-dried flour, and medua-
are mainly galactose-diglyceride (Gil and Buitrago 2002). The me-mbong. Riboflavin is well retained in boiled cassava, gari,
predominant fatty acids are palmitate and oleate (Hudson and and smoked-dried cassava flour obtained after retting of cassava
Ogunsua 1974). The protein content is low at 1% to 3% on a DM root with peel. In contrast, the losses of vitamin B 2 (riboflavin)
182     COMPREHENSIVE REVIEWS IN FOOD SCIENCE AND FOOD SAFETY—Vol. 8, 2009
Nutritional value of cassava . . .
Figure 1 --- Cassava utilization (A) and consumption per day (B) throughout the world (produced by the Intl. Inst. for
Tropical Agriculture Geospatial lab [Ibadan, Nigeria]) based on FAOSTAT 2003 data at http://faostat.fao.org/site/609/
default.aspx.
Table 3 --- Nutritional composition of different kinds of foods (100 g) for comparison to cassava root.a
                                      Water         Energy           Energy           Protein           Total          Ash         Carbohydrate by                Dietary          Sugars
Food                                   (g)           (kcal)           (kj)              (g)           lipid (g)        (g)          difference (g)                fiber (g)          (g)
Cassava, raw root                     59.68            160              667                1.36         0.28           0.62                38.06                     1.8             1.7
Potato, raw                           79.34             77              321                2.02         0.09           1.08                17.47                     2.2             0.78
Cereals
Wheat flour, unenriched               11.92            364             1523            10.33            0.98           0.47                76.31                     2.7             0.27
Bread, wheat                          35.74            266             1115            10.91            3.64           2.2                 47.51                     3.6             5.75
Rice, white, unenriched               12.89            360             1506             6.61            0.58           0.58                79.34                      ---             ---
Corn, sweet, white, raw               75.96             86              358             3.22            1.18           0.62                19.02                     2.7             3.22
Corn, yellow                          10.37            365             1527             9.42            4.74           1.2                 74.26                     7.3             0.64
Sorghum                                9.2             339             1418            11.3             3.3            1.57                74.63                     6.3              ---
Vegetables (raw)
Green beans                           90.27             31              129             1.82            0.12           0.66                 7.13                     3.4             1.4
Carrots                               88.29             41              173             0.93            0.24           0.97                 9.58                     2.8             4.74
Spinach                               94                14               59             1.5             0.2            1.8                  2.5                       ---             ---
Lettuce, green leaf                   95.07             15               61             1.36            0.15           0.62                 2.79                     1.3             0.78
Soybeans, green                       67.5             147              614            12.95            6.8            1.7                 11.05                     4.2              ---
Animal products
Raw egg (white)                       87.57             52              216            10.9             0.17           0.63                  0.73                    0               0.71
Cheese, Cheddar                       36.75            403             1684            24.9            33.14           3.93                  1.28                    0               0.52
Milk (whole)                          88.32             60              252             3.22            3.25           0.69                  4.52                    0               5.26
Raw fish (trout)                      71.42            148              619            20.77            6.61           1.17                  0                       0                0
a All values were obtained from the USDA Natl. Nutrient database for standard references (http://www.nal.usda.gov/fnic/foodcomp/search/). Nutrient values and weights are for the edible
portion.
Table 4 --- Amino acid profile of cassava.a                                                     comparison with liver (121 mg/kg FW) and egg yolk (58.7 mg/kg
                                                                                                FW), although the iron from plant origin is generally less bioavail-
                            Content in roots                   Content in leaves
                                                                                                able than iron from animal food sources. Iron and zinc content
                     % wet % dry    %                     % wet % dry    %                      in CLM are comparable to those reported for sweet potato leaves
Amino acid            wt    wt   proteinb                  wt    wt   proteinb                  and peanut leaves (Table 9). Calcium content is comparable to
                                                                                                those of peanut and broccoli, and magnesium content surpasses
Arginine              0.10          0.29      11.0          0.30       1.48         5.30        that of broccoli but is below those of peanut and sweet potato.
Histidine             0.02          0.07       2.60         0.13       0.66         2.30        Thus, mineral content of CLM is comparable with that of other
Isoleucine            0.01          0.03       1.00         0.33       1.67         5.90        leaves (Wobeto and others 2006).
Leucine               0.11          0.31      11.70         0.54       2.72         9.70           The vitamin content of cassava leaves (Table 2) is richer in
Lysine                0.02          0.07       2.60         0.37       1.87         6.70        thiamin (vitamin B 1 , 0.25 mg/100 g) than legumes and leafy
Methionine            0.01          0.03       1.00         0.07       0.36         1.30        legumes, except for soybeans (0.435 mg/100 g). The leaves have
Phenylalanine         0.01          0.03       1.00         0.18       0.92         3.30        more thiamin than several animal foods including fresh egg,
Threonine             0.01          0.03       1.00         0.27       1.35         4.80        cheese, and 3.25% fat whole milk (Table 6). The riboflavin (vi-
Tryptophan              –           0.29       0.50         0.05       0.24         0.80        tamin B 2 ) content of cassava leaves (0.60 mg/100 g) surpasses
Valine                0.01          0.04       1.50         0.20       0.99         3.50        that of legumes, leafy legumes, soybean, cereal, egg, milk, and
Alanine               0.05          0.15       5.70         0.34       1.70         6.10        cheese (Table 6). The niacin content (2.4 mg/100 g) is comparable
Aspartic acid         0.04          0.13       4.90         0.49       2.44         8.70        to that of maize (2 mg/100 g), and surpasses those reported for
Cysteine              0.003         0.01       0.40         0.04       0.21         0.70        legumes and leafy legumes, milk, and egg (Table 6). The vitamin
Glutamic acid         0.05          0.15       5.70         0.40       1.99         7.10        A content of cassava leaves is comparable with that of carrots
Glycine               0.003         0.01       0.40         0.35       1.73         6.20        and surpasses those reported for legumes and leafy legumes. The
Proline               0.01          0.03       1.00         0.18       0.88         3.10        vitamin C content (60 to 370 mg/100 g) of cassava leaves is high
Serine                0.01          0.04       1.50         0.34       1.68         6.00        compared to values reported for other vegetables (Table 6). Thus,
Tyrosine              0.003         0.01       0.40         0.18       0.89         3.20        the overall vitamin content of the leaves is comparable and in
a Adapted from Gil and Buitrago   (2002).                                                       certain cases better than those reported for most legumes, leafy
b Content of total protein (%).
                                                                                                legumes, cereals, egg, milk, and cheese.
                                                                                                Fiber
reported: from 61.5 to 270 mg iron/kg DM, 30 to 63.7 mg zinc/kg                                    The fiber content of cassava leaves is high (Table 2) compared
DM, 50.3 to 263 mg manganese/kg DM, 6.2 to 50 mg copper/kg                                      to the fiber content of legumes and leafy legumes reported in
DM, 2.3 to 3 g sulfur/kg DM, 2.6 to 9.7 g magnesium/kg DM,                                      Table 3 and ranges between 1 and 10 g/100 g FW. Dietary fiber
0.4 to 16.3 g calcium/kg DM, and 8 to 16.9 g potassium/kg                                       is considered part of a healthy diet and can reduce problems of
DM (Barrios and Bressani 1967; Gomez and Valdivieso 1985;                                       constipation. Although recent evidence is mixed, fiber may help
Nwokolo 1987; Ravindran and others 1992; Aletor and Adeo-                                       prevent colon cancer (Rock 2007). The rich fiber of cassava may
gun 1995; Awoyinka and others 1995; Chavez and others 2000;                                     assist intestinal peristalsis and bolus progression (Favier 1977),
Madruga and Câmara 2000). Cassava leaf meal is rich in iron in                                 but if fiber content from any source is too high, it will have
Table 5 --- Mineral content of 100 g of various foods for comparison to cassava root.a
                                            Cab          Fe              Mg             P              K              Na             Zn               Cu               Mn           Se
Food                                        mg           mg              mg             mg             mg             mg             mg               mg               mg           μg
Cassava, raw root            20.6         0.087           0.048         0.854           0.107           0.088          27         0                1            0        0.19           1.9
Potato, raw                  19.7         0.08            0.032         1.054           0.296           0.295          16         0                0            0        0.01           1.9
Cereals
Wheat flour,                   0          0.12            0.04          1.25            0.438           0.044          26         0                0            0        0.06           0.3
  unenriched
Bread, wheat                   0.2        0.373           0.311         5.19            0.82            0.119          85         0                0            0        0.19           4.9
Rice, white,                   0          0.07            0.048         1.6             1.342           0.145           9         0                0            0         ---          ---
  unenriched
Corn, sweet,                   6.8        0.2             0.06          1.7             0.76            0.055          46         0                0            0        0.07           0.3
  white, raw
Corn, yellow                   0          0.385           0.201         3.627           0.424           0.622           19        0              11             0        0.49           0.3
Sorghum                        0          0.237           0.142         2.927             ---             ---          ---        0               0             0         ---          ---
Vegetables (raw)
Green beans                  16.3         0.084           0.105         0.752           0.094           0.074          37         0              35             0        0.41          14.4
Carrots                       5.9         0.066           0.058         0.983           0.273           0.138          19         0             841             0        0.66          13.2
Spinach                      30           0.04            0.13          0.5             0.312           0.304          15         0             220             0         ---           ---
Lettuce, green leaf          18           0.07            0.08          0.375           0.134           0.09           38         0             370             0        0.29         173.6
Soybeans, green              29           0.435           0.175         1.65            0.147           0.065         165         0               9             0         ---           ---
Animal products
Raw egg (white)                0          0.004           0.439         0.105           0.19            0.005           4         0.09            0            0         0              0
Cheese, Cheddar                0          0.027           0.375         0.08            0.413           0.074          18         0.83          265          258         0.29           2.8
Milk, whole                    0          0.044           0.183         0.107           0.362           0.036           5         0.44           28           28         0.06           0.2
Raw fish (trout)               0.5        0.35            0.33          4.5             1.94            0.2            13         7.79           17           17         0.2            0.1
a All values were obtained from the USDA Natl. Nutrient database for standard references (http://www.nal.usda.gov/fnic/foodcomp/search/). Nutrient values and weights are for the
edible portion.
b Vitamin C = total ascorbic acid; Vitamin A is represented as retinol activity equivalents (RAE) and retinol refers to vitamin A in the preform; vitamin E = α-tocopherol; vitamin
K = phylloquinone.
Wet root (g)                       100           77.0         87.6        49.2                38.5             25.3 to 29.6             27.9 to 34.0     66.8
Dry matter (g)                      40.0         32.3         28.3        21.6                29.7             21.3 to 25.6             20.8 to 28.7     19.0
Calories                           157          127          112          86                 119                85 to 102                 83 to 115      76
Protein (g)                          1.0          0.48         0.38        0.18                0.37            0.16 to 0.22             0.26 to 0.51      0.16
Fat (g)                              0.1          0.1          0.04        0.02                0.2             0.04 to 0.06             0.04 to 0.12      0.03
Carbohydrates (g)                   37.9         31.0         27.4        21.2                28.8             20.9 to 25.1             20.3 to 28.1     18.8
Fiber (g)                            1.3          0.6          0.5         0.4                 0.6                  0.4                  0.3 to 0.6       0.3
Ash (g)                              0.90         0.57         0.46        0.21                0.34            0.16 to 0.19             0.24 to 0.50      0.06
Calcium (mg)                        26           13           12           7                  10                6.0 to 8.0              7.0 to 15.0      11
Phosphorus (mg)                     47           39           31          13                  18               9.0 to 11.0              10.0 to 21.0      7
Iron (mg)                            3.5          0.4          0.4         3.1                 1.5              0.2 to 0.7               0.8 to 11.9      0.2
Thiamin (μg)                        72           31           20          10                  18                6.0 to 12.0                  13           3
Riboflavin (μg)                     34           18           16          21                  15               10.0 to 12.0              8.0 to 21.0      6
Niacin (mg)                          0.73         0.52         0.41        0.16                0.33            0.11 to 0.18             0.17 to 0.37      0.03
Vitamin C (mg)                      33           20            1           1                   2                     0                        0           0
a Adapted   from Favier (1977).
Table 8 --- Comparison of the amino acid profile of cassava Table 9 --- Comparison of sweet potato leaf and peanut
with egg, on a 16 g N basis (adapted from Favier 1977).     leaf nutrients with cassava leaf meal (CLM).a
                                               Cassava leavesb                  Nutrient (100 g dry weight)                   CLM        Sweet potato   Peanut
                             Egg                                      Cassava
Amino acid                 proteina         Jamaica        Brazil      rootc    Protein (g)                                    28.1           30.6        26.6
                                                                                β-Carotene (mg)                                88.0           75         113.3
                                            % on a basis of 16 g N
                                                                                Vitamin C (mg)                                 90.2          141.7       293.3
Tryptophan                       1.5          1.5            2.1         –
                                                                                Iron (mg)                                      16.7           14.7        16
Threonine                        4.9          2.8            4.9        4.7
                                                                                Zinc (mg)                                       5.08           3.33         5.33
Isoleucine                       8.0          5.0            4.8        1.8
                                                                                Manganese (mg)                                 14.1            ---         ---
Leucine                          9.2          8.9            8.8        2.9
                                                                                Magnesium (mg)                                229.3          493.3       676.7
Lysine                           3.9          7.2            6.3        7.2     Calcium (mg)                                 1509.4          623.3      1236.7
Methionine                       4.1          1.7            1.7        1.0
                                                                                a Adapted   from Wobeto and others (2006).
Cysteine                         2.4          1.4            1.0         –
Phenylalanine                    6.3          5.8            5.5        2.1
Tyrosine                         4.5          4.2            3.9        1.6     have important deficits in cysteine and tryptophan, and have rel-
Valine                           7.3          5.8            5.6        2.6     atively low concentrations of isoleucine, leucine, phenylalanine,
Arginine                         6.4          5.3            6.1       14.9     tyrosine, and valine (Jacquot 1957). The leaves have tryptophan
Histidine                        2.1          2.2            2.6        1.7     concentrations comparable to those found in eggs, but the cys-
a   Mitchell and Block (1946).                                                  teine content in leaves is only about half that of the egg (Jacquot
b
c
    Rogers and Milner (1963).                                                   1957).
    Busson (1965).
                                                                                   Cassava roots have a large excess of arginine, while the arginine
negative effects in humans. Fiber can be a nutritional concern                  content of leaves is relatively low and comparable to that of
because it can decrease nutrient absorption in the body (Baer                   eggs. Both leaves and roots have an excess of lysine, which is
and others 1996). Excess fiber will increase fecal nitrogen, cause              approximately twice as high as what has been observed in eggs
intestinal irritation, and reduce nutrient digestibility, in particular         (Jacquot 1957) on a basis of 16 g nitrogen. Although cassava
protein digestibility (Favier 1977; Baer and others 1996). It is                roots do not have a well-balanced amino acid profile and are of
important to optimize the utilization of nutrients from cassava                 lower nutritional value because of low protein quantity, cassava
because nutrient deficiencies are more prevalent in regions where               leaves have good protein content. Therefore, cassava leaf proteins
cassava is used as a staple food.                                               could be used to improve the nutritional value of a diet primarily
                                                                                made up of cassava roots. However, methionine and cysteine
                                                                                would still be limiting amino acids (Gil and Buitrago 2002), so a
Comparison of Nutritional Value of Cassava Roots                                methionine and cysteine source should be added to the diet to
and Cassava Leaves                                                              supplement these 2 sulfur-containing amino acids.
   Table 2 compares the proximate, vitamin, and mineral compo-                     The lipid content is 10 times higher in leaves than in roots. Al-
sitions of cassava leaves and roots. The roots are twice as rich as             though the lipids and lipid-soluble components such as chloro-
the leaves in carbohydrates, but the leaves contain more protein,               phyll, resin, and xanthophylls are much more concentrated in
lipid, minerals, vitamins, and fiber. The total protein content in              leaves, some of them, such as volatile fatty acids, chlorophyll,
cassava leaves is 5 to 10 times higher than in roots and is com-                and resin, do not bring significant energy to the diet. Therefore,
parable with the protein content of egg (Jacquot 1957) based on                 the energy density of the lipid is lower in leaves than in roots (Gil
grams of nitrogen. The protein content of cassava leaves is sim-                and Buitrago 2002).
ilar to those of sweet potato leaves and peanut leaves (Table 9)                   The mineral content of cassava leaves is 2 to 5 times higher
(Wobeto and others 2006). There is a significant deficit in me-                 than that of the roots. The roots typically have more phospho-
thionine for both cassava leaves and roots. Cassava roots also                  rus, but the leaves have a greater concentration of calcium (Gil
                                                           Vol. 8, 2009—COMPREHENSIVE REVIEWS IN FOOD SCIENCE AND FOOD SAFETY                               187
CRFSFS: Comprehensive Reviews in Food Science and Food Safety
and Buitrago 2002). The calcium content in the leaves is 100             ties. Indeed, phytic acids can reduce free ion radical generation
times higher than in roots and the phosphorus content is 2 to 3          and thus peroxidation of membranes by complexing iron, and
times higher in roots than in the leaves. Cassava leaves are more        phytate may protect against colon cancer (Graf and others 1987).
concentrated than the roots in vitamins and the minerals iron,           Phytate was able to reduce serum cholesterol and triglycerides in
potassium, magnesium, copper, zinc, and manganese. Indeed,               an animal model fed a cholesterol-enriched diet (Jariwalla 1999).
thiamin and niacin contents are 4 to 5 times higher in leaves than       Dephosphorylation of phytate occurs during processing of cas-
in roots, and riboflavin and vitamin C are 10 to 12 times higher         sava, especially during fermentation when > 85% of phytate is
in the leaves. Cassava leaves have a high quantity of vitamin A in       removed (Marfo and others 1990). The possibility of genetic mod-
the form of provitamin A carotenoids. Vitamin E, however, is low         ification of cassava to reduce phytate concentrations has not yet
in both the leaves and roots (Gil and Buitrago 2002).                    been investigated (Montagnac and others 2009).
   Thus, cassava roots are of lower nutritional value regarding             The polyphenol content (tannins) in cassava leaves is increased
mineral, vitamin, lipid, and protein contents, but the leaves are        with the maturity of the plant. Cassava leaf meal has a polyphe-
well provided in these, and should be added to a diet consisting         nol content of 2.1 to 120 mg/100 g DM (Wobeto and others
mainly of roots. Cassava leaves are typically served boiled and          2007). Polyphenols can form insoluble complexes with divalent
mixed with other vegetables, such as okra and beans. The fiber           ions such as iron, zinc, and copper. They can also inactivate
content of leaves is greater than that of roots (3.5 times more), and    thiamin, bind certain salivary and digestive enzymes, and en-
so, it may limit the absorption of minerals, vitamins, and proteins      hance secretion of endogenous protein. Consequently, they in-
by the body and may be a restricting nutritional factor.                 hibit nonheme-Fe absorption, reduce thiamin absorption and the
                                                                         digestibility of starch, protein (Silva and Silva 1999), and lipids,
                                                                         and also interfere with protein digestibility (Bravo 1998). How-
Cassava and Selected Antinutrients                                       ever, tannins also have antioxidant and anticarcinogen proper-
   Analyzing the nutritional value of cassava, it appears that cas-      ties that can benefit humans (Chung and others 1998; Chen and
sava roots are a good carbohydrate source and cassava leaves are         Chung 2000; Alessio and others 2002; Matuschek and Svanberg
good mineral, vitamin, and fiber sources for humans. However,            2002; Nakagawa and others 2002). Catechins (catechin, cate-
cassava contains antinutrients and toxic substances that interfere       chin gallate, gallocatechin) and flavone 3-glycosides (rutin and
with the digestibility and the uptake of some nutrients. Neverthe-       kaempferol 3-rutinoside), suggested to have cardiovascular health
less, depending on the amount consumed, these substances can             benefits, have been identified in cassava roots (Buschmann and
also bring benefits to humans.                                           others 2000). Anthocyanidins (cyanidin and delphinidin) have
   Cyanide is the most toxic factor restricting the consumption of       been identified in cassava leaves (Reed and others 1982). Pro-
cassava roots and leaves. Indeed, cassava, particularly its bitter va-   cessing cassava leaves reduces the polyphenol content, but 50%
rieties, has a cyanide level higher than the FAO/WHO (1991) rec-         to 60% is retained; again, it has been suggested that genetic
ommendations, which is < 10 mg cyanide equivalents/kg DM, to             modifications might well reduce total polyphenol content (Fasuyi
prevent acute toxicity in humans. Cassava leaves have a cyanide          2005).
content ranging from 53 to 1,300 mg cyanide equivalents/kg of               Oxalates are antinutrients affecting calcium and magnesium
DW (Siritunga and Sayre 2003; Wobeto and others 2007), and               bioavailability (Massey 2007) and form complexes with proteins,
cassava root parenchyma has a range of 10 to 500 mg cyanide              which inhibit peptic digestion (Oboh 1986). Oxalate content
equivalents/kg DM (Arguedas and Cooke 1982; Dufour 1988;                 ranges from 1.35 to 2.88 g/100 g DM for cassava leaf meal (Fon-
Siritunga and Sayre 2003); both of these are much higher than            seca 1996; Corrêa 2000; Wobeto and others 2007). The negative
what is recommended. Several health disorders and diseases have          effect of oxalates on humans depends on the level of both oxalate
been reported in cassava-eating populations. Consumption of 50           and calcium in the cassava leaves. Wobeto and others (2007) re-
to 100 mg of cyanide has been associated with acute poisoning            ported that the calcium-to-oxalate ratio of 5 cassava cultivars
and has been reported to be lethal in adults (Halstrom and Moller        was greater than 0.44%, which means that oxalate levels found
1945). The consumption of lower cyanide amounts are not lethal           in cassava leaf meal do not diminish the uptake of calcium.
but long-term intake could cause severe health problems such                Saponins are plant glycosides that are studied for their potential
as tropical neuropathy (Osuntokun 1994), glucose intolerance,            health benefits, particularly those derived from ginseng and soy
konzo (spastic paraparesis) (Ernesto and others 2002), and, when         (Bachran and others 2008; Yin and others 2008). Saponins are
combined with low iodine intake, goiter and cretinism (Delange           considered the bioactive component of ginseng responsible for
and others 1994).                                                        its metabolic and potential health effects (Yin and others 2008;
   In addition, the nitrate content in cassava leaves ranges from        Christensen 2009). They have antitumor properties and may of-
43 to 310 mg/100 g DM (Corrêa 2000; Wobeto and others 2007).            fer synergistic effects when used in combination with drug ther-
Cassava-eating populations ingesting cyanide and high amounts            apy (Bachran and others 2008). Saponins can also act on the
of nitrates and nitrites have the risk of developing stomach cancer.     central nervous system of humans with potential therapeutic ef-
Cassava-eating individuals tend to have a high amount of thio-           fects (Nah and others 2007). Cassava leaf meal has a steroidal
cyanate in the stomach due to cyanide detoxification by the body,        saponin content ranging from 1.74 to 4.73 g/100 g DM (Wobeto
which may catalyze the formation of carcinogenic nitrosamines            and others 2007), which compares to those found for soybeans
(Mirvish 1983; Maduagwu and Umoh 1988; Onyesom and Okoh                  (0.07 to 5.1 g/100 g DM) (Fenwick and Oakenfull 1983; Ireland
2006).                                                                   and Dziedzic 1985; Schiraiwa and others 1991), but is lower
   Phytate (inositol hexakisphosphate) is another compound               than those observed in alfalfa (5.6 g/100 g DM) and beet leaves
found in high abundance in cassava, with approximately                   (5.8 g/100 g DM) (Fenwick and Oakenfull 1983). Saponin con-
624 mg/100 g in roots (Marfo and others 1990). Phytic acid is            tent increases in cassava leaf meal with plant maturity (Wobeto
able to bind cations such as magnesium, calcium, iron, zinc, and         and others 2007).
molybdenum and can, therefore, interfere with mineral absorp-               Cassava leaf meal has a trypsin inhibitor content of 3.79 in-
tion and utilization which may affect requirements (Hambidge             hibited trypsin unit (ITU)/mg DM at the starch accumulation
and others 2008). It may also bind proteins preventing their com-        phase (17-mo-old plant) and of 11.14 ITU/mg DM at the leaf
plete enzymatic digestion (Singh and Krikorian 1982). However,           development phase (12-mo-old plant) (Corrêa and others 2004).
phytic acids also have antioxidant and anticarcinogenic proper-          Trypsin inhibitor has adverse effects on the pancreas. Indeed,
188   COMPREHENSIVE REVIEWS IN FOOD SCIENCE AND FOOD SAFETY—Vol. 8, 2009
Nutritional value of cassava . . .
Liener (1977) has demonstrated that for species with a pancreas       Moreover, this cassava hybrid also had richer protein content
comprising more than 0.3% of the body weight, trypsin inhibitor       in the peel (8.06%) than typical cassava cultivars (from 1.11% to
feeding in these species will produce an enlargement of the pan-      2.09%) (Nassar and Dorea 1982). Further research has continued
creas. Unheated soybean trypsin inhibitor decreases the activity      to indicate the feasibility of selecting interspecific hybrids that are
of rat, monkey, human, bovine, porcine, and mink trypsins at a        rich in both crude protein and amino acids to improve the protein
rate of 90% to 100% and rat, monkey, and human total prote-           value (Nassar and Souza 2007). This interspecific cassava hybrid
olytic activity by up to 40% (Struthers and Macdonald 1983).          has an improved amino acid profile with 10 times more lysine
However, protease inhibitors may suppress carcinogenesis (Park        and 3 times more methionine than the common cultivar.
and others 2007).                                                        Transgenic approaches to reduce cyanogen in cassava have
                                                                      focused on suppressing cyanogen synthesis or accelerating
                                                                      cyanogen breakdown (Siritunga and Sayre 2007). One poten-
Biofortification and Processing Methods to Improve the tial benefit of lowering cyanogen content is the facilitation of free
Nutritional Value of Cassava                                          cyanide assimilation into amino acids (Siritunga and Sayre 2007).
   Cassava is a target for biofortification because of its impor- Thus, reducing toxic cyanogens would have the added benefit of
tance as a staple crop. The Bill and Melinda Gates Foundation improving the protein value of the roots.
has supported a global effort to develop cassava germplasm en-
riched with bioavailable nutrients since 2005 (BioCassava Plus Postharvest processing to enhance protein
[http://biocassavaplus.org/]). This initiative is called BioCassava      Another approach that increases the protein content and qual-
Plus and has 6 major objectives: to increase the minerals zinc ity of ready-to-eat cassava products is the development of posthar-
and iron, increase protein, increase vitamins A and E, decrease vest processing techniques. Crude protein of cassava root and
cyanogen content, delay postharvest deterioration, and develop leaf by-products can be increased by solid-state fermentation via
virus-resistant varieties.                                            Aspergillus niger (Iyayi and Losel 2001), while also decreasing
                                                                      cyanogen content by up to 95% (Birk and others 1996). Smith and
Biofortified cassava and protein value                                others (1986) reported a significant increase in the protein con-
   Cassava roots, with a crude protein content of about 1.5%, are tent of cassava roots by solid-state fermentation via Sporotrichum
low in protein and some essential amino acids. To date, several pulverulentum. This fungus was able to produce 30.4 g of high-
                                                                                                                                            ◦
different strategies have been investigated to improve the protein quality protein per 100 g of dry cassava in 48 h at 45 C. The
content and the amino acid composition of cassava ready-to-           protein  bioavailability  of  fermented       cassava  leaves  was  similar to
eat products. To engineer improved storage proteins with bal- that of soybean pressed cake diets delivered to ruminants, and,
anced amino acid composition in cassava tubers, Zhang and therefore, fermented cassava leaves can replace soybean as a
others (2003a) successfully transferred a 284 bp synthetic gene source of protein (Bakrie and others 1996).
(ASP1) coding for a 11.2 kDa-storage protein rich in essential           The development of cassava leaf protein concentrates with low
amino acids (80%) into embryonic suspensions of cassava using fiber could enhance the protein value of cassava meals. Crude
Agrobacterium. They observed stable integration and expression protein content of cassava leaf protein concentrate is twice (42%
of ASP1 in cassava leaves and primary roots. However, contrary to 43%) that of cassava leaf meal (22%) (Castellanos and others
to the results obtained for transgenic sweet potatoes (Egnin and 1994). Indeed, Fasuyi and Aletor (2005) reported that cassava
others 2001) and tobacco lines (Kim and others 1992) where leaf protein concentrates had crude protein, fat, and gross energy
ASP1 had been over-expressed, no significant differences in pro- content higher than those of cassava leaf meals and lower crude
tein content and in the overall amino acid composition of cassava fiber and ash contents. Table 10 shows the proximate composi-
leaves were observed between transgenic and wild-type cassava tion of cassava leaf meal and cassava leaf protein concentrate.
lines (Zhang and others 2003a). Nonetheless, analysis of 1-y-old Although methionine (2.48 g/16 g N) and cysteine were limited
cassava plants from the 2nd vegetative generation grown under in concentrates, the amino acid analysis determined that lysine
greenhouse conditions showed an increase in protein content (6.80 g/16 g N), leucine (9.65 g/16 g N), valine (6.30 g/16 g N),
and essential amino acid composition in storage roots of sev- and tryptophan (2.31 g/16 g N) exceeded those of soybean, fish,
eral transgenic lines (Zhang and others 2004). Two cassava root and egg reported by the FAO/WHO (1973). In addition, the wa-
specific promoters related to vascular expression and secondary ter absorption capacity (181.5% ± 45.4%), the fat absorption
growth were identified (Zhang and others 2003b), which repre-
sent valuable candidates for targetting the protein ASP1 in storage
roots for genetic improvement. Currently, studies to improve the Table 10 --- Comparison of cassava leaf protein concen-                    a
levels of expression and accumulation of the ASP1 protein into trate and cassava leaf meal on a nutritional basis.
cassava tubers are underway (Zhang and others 2004).                                                                           Variety
   Researchers have also tried to improve the nutritional value
of cassava by crossbreeding wild-type varieties. Two hybrids                                                                         Cassava leaf
showed promising results regarding protein content compared                                                    Cassava leaf             protein
to typical cassava cultivars. The interspecific hybrid of cassava                                                    meala          concentratesb
(UnB 033) and Manihot dichotoma showed higher protein con- Proximate composition                                  1         2         1         2
tent (26.4%) in its leaves compared to cassava cultivars (24.25%)
(Nassar and others 2004). This hybrid also resulted in a 5-times Crude protein (g/kg DM)                     343.0      363.0      500.0      493.0
higher content of manganese and zinc than those of typical cas- Crude fiber (g/kg DM)                        127.0      115.0       18.0        14.0
sava cultivars (EB01). The leaf cyanide content was moderate, that Ether extract (g/kg DM)                     75.0      70.0      216.0      224.0
is, 128.5 ± 11.7 mg cyanide/kg FW (Nassar and others 2004). A Ash (g/kg DM)                                    72.0      69.0       93.0        79.0
2nd valuable hybrid was achieved by crossing with Manihot oli- Nitrogen free extract (g/kg DM) 383.0 382.0 115.0                              141.0
gantha (Nassar and Dorea 1982). Protein content in the roots was Gross energy (MJ/kg)                          47.1      47.2       53.6        50.1
twice that of typical cassava cultivars. For peeled tuber, the inter- Digestible energy (MJ/kg)                  ---       ---      46.7        47.6
specific hybrid had a protein content of 4.5%, while the protein a Adapted from Fasuyi (2005).
content observed in cassava cultivars ranged from 0.9% to 1.4%. b Adapted from Fasuyi and Aletor (2005); Fasuyi (2006).
                                                  Vol. 8, 2009—COMPREHENSIVE REVIEWS IN FOOD SCIENCE AND FOOD SAFETY                            189
CRFSFS: Comprehensive Reviews in Food Science and Food Safety
capacity (19.2% ± 1.2% to 40.8% ± 1%), the emulsion capacity             be increased with addition of a modified bacterial AGPase. This
(32.5% ± 8.3%) and stability (42.9% ± 2.9%), the least gelation          would theoretically enhance cassava root starch production. Un-
concentration (12.5% ± 3.4%), the foaming capacity (32.1% ±              der greenhouse conditions, Ihemere and others (2006) observed
7.7%) and stability (10.2 ± 4.1 cm3 ), and the solubility of cas-        an AGPase activity 70% higher than that of wild types (pH 7.5,
sava leaf protein concentrates in acid and alkaline media support        25 ◦ C) and a 2.6-fold increase in total tuberous root biomass
the nutritive potential of cassava leaf protein concentrate (Fasuyi      for transgenic cassava plants. Although the density of starch was
and Aletor 2005). These properties of cassava leaf protein con-          not modified between the transgenic and wild-type lines, an in-
centrate allow formulation into viscous products such as soups,          crease in tuberous root size and number (biomass) resulted in an
protein-rich carbonated beverages, and curds; or as additives for        increase in total starch.
gel formation in food products. They also show the ability of cas-
sava leaf protein concentrate to form and stabilize emulsions in         Postharvest technique to enhance energy density
food products (Fasuyi and Aletor 2005). Therefore, cassava leaf             Another strategy to enhance the energy density of cassava by-
protein concentrate is a good alternative protein for human and          products is to add amylase. In the case of weaning food products,
animal nutrition, but due to the few limiting amino acids it should      Treche and others (1994) demonstrated that the energy density
not be the exclusive source of protein. Growth bioassays in rats         of cassava gruels would double, and would maintain an ac-
showed that cassava leaf protein concentrate should be fed along         ceptable consistency when plant amylase sources such as flour
with another viable protein source, because the concentrate with         from malted cereals are added. The alpha-amylases hydrolyze
and without dl-methionine supplements did not support growth             the starchy component of cassava gruel into maltose and dex-
(Fasuyi 2005).                                                           trins of low molecular weight and of low water-binding capac-
    Regarding techniques to produce protein concentrates, ultra-         ity, which allows a reduction in the viscosity of cassava liquid
filtration is more efficient than that of acidic thermocoagulation       gruel.
(Table 11) (Castellanos and others 1994). Indeed, ultrafiltration
provides a better protein efficiency ratio (1.81), protein digestibil-   Biofortified cassava and vitamin A
ity in vitro (85%), and the availability of lysine (90%). It also in-       Vitamin A is a fat-soluble vitamin playing an important role
creases the amino acid content, especially of lysine (5.05 g/100 g       in vision, bone growth, reproduction, and in the maintenance of
protein), threonine (4.15 g/100 g protein), tryptophan (0.7 g/100 g      healthy skin, hair, and mucous membranes (FAO/WHO 2002).
protein), and sulfur-containing amino acids. However, sulfur-            Identified as a widespread public health problem in 37 countries
containing amino acids and tryptophan are still deficient after          worldwide, vitamin A deficiency is the most common cause of
processing compared with FAO’s recommendations.                          childhood blindness. It is estimated that 228 million children are
                                                                         affected and 500000 children become partially or totally blind
Biofortified cassava and energy density                                  every year as a result of vitamin A deficiency (WHO/FAO 2003).
   Although cassava root has one of the highest rates of CO 2            The geographical areas most affected by vitamin A deficiency
fixation (43 μmol CO 2 /m2 /s) and of sucrose synthesis for a C3         are tropical areas where cassava is a staple crop, for example,
plant, carbohydrate yield is below its full potential. Therefore, it     Brazil, Africa, and Asia (Shrimpton 1989). Biofortification of sta-
has been hypothesized that cassava plants can be engineered to           ple crops with provitamin A carotenoids is an emerging strategy
enhance their starch yield. The catalyst of the first dedicated, rate-   to address the vitamin A status of the poor (Tanumihardjo 2008;
limiting step of starch synthesis is ADP-glucose pyrophosphory-          Tanumihardjo and others 2008).
lase (AGPase). Its activity is a key factor for the enhancement             Cassava root contains small amounts of β-carotene, a provita-
of starch production. Muller-Rober and others (1992) observed            min A carotenoid, which can be converted as needed into retinal,
severely reduced starch production and accumulation of sucrose           reduced to retinol, and stored in the liver esterified to fatty acids.
and glucose in potato tubers as a result of inhibiting AGPase ex-        The bioconversion of β-carotene to vitamin A in the body is nat-
pression. Because bacterial AGPase activity is several hundred-          urally regulated and therefore β-carotene has little potential for
folds higher than that of the cassava plant, Ihemere and others          toxicity compared with high intake of vitamin A-fortified foods
(2006) hypothesized that AGPase activity of cassava roots could          (Tanumihardjo 2008). Fortified foods, such as sugar in Guatemala
Table 11 --- Comparison of ultrafiltration and thermocoagulation processes on nutritional value of cassava leaf protein
concentrates.a
                                                    Casein                Cassava                                              Acidic
Proximate analysis                                 reference                leaf                Ultrafiltration           thermocoagulation
(Dary and Mora 2002) and Nicaragua (Ribaya-Mercado and oth-           cient at quenching the destructive potential of singlet oxygen (Di
ers 2004), and supplements contain highly bioavailable pre-           Mascio and others 1989). Red cassava with substantial lycopene
formed vitamin A and uptake is not regulated like bioconversion.      is currently being distributed to small-scale farmers throughout
Utilizing biofortified cassava with enhanced β-carotene (Figure 3)    Brazil (Nassar 2007). Lutein and zeaxanthin might act as antioxi-
would be a sustainable strategy to reduce the prevalence of vi-       dants in the macular region of the human retina (Snodderly 1995;
tamin A deficiency in areas where cassava is a staple food. In        Pauleikhoff and others 2001). The possible role of lutein in pre-
addition, β-carotene can act as an antioxidant to protect cells       venting age-related eye disease is currently under investigation.
and tissues from the damaging effects of free radicals and singlet       Screening, selecting, and crossbreeding varieties of cassava
oxygen species (Paiva and Russell 1999). For example, in Mongo-       with high content of carotene is currently underway and shows
lian gerbils fed biofortified carrots or vitamin A supplements, the   potential as a dietary source (Nassar and others 2005). A broad
antioxidant status of the carrot groups was higher than the sup-      distribution of carotene concentrations in cassava leaves and
plement group (Mills and others 2008). However, β-carotene’s          roots has been observed. Carotene content is 100 times higher
role in cancer prevention is not completely understood; smokers       in cassava leaves (ranging from 12 to 97 mg/100 g FW) than
taking daily β-carotene supplements had a greater risk of lung        in roots (ranging from 0.102 to 1.069 mg/100 g FW) (Iglesias
cancer and cardiovascular disease than those not taking supple-       and others 1997; Chavez and others 2003). The color intensity
ments (Goodman and others 2004).                                      of the cassava root and the carotene concentration are posi-
   Cassava also contains other interesting carotenoids that are not   tively correlated (Iglesias and others 1997). The carotene con-
provitamin A, such as the carotene lycopene and the xanthophylls      centration was 0.13 mg/100 g in white cassava roots, and 0.39
lutein and zeaxanthin. Lycopene appears to be particularly effi-      mg/100 g, 0.58 mg/100 g, 0.85 mg/100 g, and 1.26 mg/100 g
                                                                      in cream, yellow, deep yellow, and orange cassava roots, respec-
                                                                      tively. Moreover, 5 orange cassava genotypes have been found
                                                                      with β-carotene contents ranging from 2.04 to 2.55 mg/100 g
                                                                      FW in the Amazonian region of Brazil and Colombia (Iglesias
                                                                      and others 1997). These results are encouraging because using
                                                                      conservative conversion factors of 12 μg β-carotene to 1 μg vi-
                                                                      tamin A proposed by the Inst. of Medicine, Food and Nutrition
                                                                      Board (2001), this level of β-carotene would result in 170 to
                                                                      210 μg vitamin A. This range of vitamin A includes the esti-
                                                                      mated average requirement for a young child 1 to 2 y old (that
                                                                      is, 210 μg vitamin A) and represents about 40% of the estimated
                                                                      average requirement for a woman of childbearing age (that is,
                                                                      485 μg vitamin A). Yellow and orange cassava roots are a viable
                                                                      alternative for delivering provitamin A carotenoids to vitamin A-
                                                                      deficient populations that consume cassava. Considering the high
                                                                      daily intakes of cassava in several African countries (FAO 2006)
                                                                      (Figure 1B), cassava biofortified with β-carotene could readily
                                                                      impact the prevalence of night blindness due to vitamin A defi-
                                                                      ciency in women if widely adopted (WHO 2008) (Table 12).
                                                                         However, the stability of β-carotene during different process-
                                                                      ing techniques needs to be further evaluated. Carotene stability is
                                                                      genotypically dependent and cassava genotypes with the highest
                                                                      carotene content in fresh roots are not the ones with the high-
                                                                      est carotene content after processing (Iglesias and others 1997).
                                                                      For example, comparing cream and yellow cassava root clones
                                                                      after processing, the yellow roots lost more β-carotene (Chavez
                                                                      and others 2004). Carotene concentration and availability is af-
                                                                      fected by heat-processing treatments. Iglesias and others (1997)
                                                                      reported that boiling, oven-drying, and sun-drying fresh cassava
                                                                      root parenchyma reduced carotene content by 34%, 44%, and
                                                                      73%, respectively. Chavez and others (2004) pointed out that
                                                                      lyophilization and production of gari from fresh cassava root re-
                                                                      duced the initial β-carotene content by 25% and 80%, respec-
                                                                      tively. Therefore, lyophilization and boiling were most effective
                                                                      at retaining β-carotene, and sun-drying and gari production the
                                                                      least.
                                                                         Processing cassava increases the cis-β-carotene isomer con-
                                                                      tent significantly (Thakkar and others 2007; Howe and others
                                                                      2009). The percentage of cis content after processing was 30% to
                                                                      52%, which was observed for 10 genotypes of processed cassava
Figure 3 --- Boiling the cream-colored biofortified cassava           (Thakkar and others 2007). The vitamin A value of cis-β-carotene
reveals the β-carotene through the deepening of color to              is generally accepted to be less than the trans isomer and is re-
yellow. The outer brown peel was removed before boil-                 ported to be 23% to 61% for 9-cis-β-carotene and 48% to 74%
ing. The underlying white peel, which is sometimes used               for 13-cis-β-carotene in gerbil and rat models (Deming and oth-
in livestock feed, was removed after freeze-drying and                ers 2002). However, the bioconversion factor for β-carotene was
before flour preparation for an animal study (Howe and                similar in Mongolian gerbils fed different percentages of cis-β-
others 2009).                                                         carotene, that is 48% and 2.5% of total β-carotene as cis from
                                              Vol. 8, 2009—COMPREHENSIVE REVIEWS IN FOOD SCIENCE AND FOOD SAFETY                     191
CRFSFS: Comprehensive Reviews in Food Science and Food Safety
Table 12 --- The impact of biofortified cassava with 2 mg β-carotene/100 g fresh weight on intake of vitamin A in
several cassava-eating African countries. Considering that the estimated average requirement of women is 485 μg/d,
biofortified cassava could positively impact the prevalence of night blindness.
                                                Prevalence of                   Consumption                  β-Carotene                 β-Carotene                  Estimated retinol
Country                                        night blindnessa                     g/db                     content mg                retainedc mg                 equivalentsd μg
Congo, Democratic Republic                       5.4% to 9.5%                          820                        16.4                        10.8                      900 to 1800
Mozambique                                       1.6% to 8.2%                          680                        13.6                         9.0                      750 to 1500
Ghana                                            4.6% to 12.8%                         600                        12                           7.9                      660 to 1300
Benin                                            4.1% to 14.7%                         430                         8.6                         5.7                      480 to 950
Guinea                                           7.9% to 21.3%                         360                         7.2                         4.8                      400 to 800
Rwanda                                           4.8% to 11.5%                         350                         7                           4.6                      380 to 770
Madagascar                                       1.7% to 15.3%                         320                         6.4                         4.2                      350 to 700
Nigeria                                          4.9% to 11.1%                         310                         6.2                         4.1                      340 to 680
a The range of prevalence in different communities of night blindness (a serious sequela of vitamin A deficiency) in women of childbearing years (WHO http://www.who.int/vmnis/vitamina/
data/database/countries/en/index.html).
b FAOSTAT agricultural data (database) Rome, Italy: FAO http://faostat.fao.org/site/609/default.aspx#ancor.
c An optimistic retention factor of 66% was used as reported for boiled cassava (Iglesias and others 1997).
d Estimated retinol equivalents were calculated using conversion factors of 6 μg β-carotene (FAO/WHO 2002) and 12 μg β-carotene to 1 μg retinol (Inst. of Medicine 2001) to encompass a
range of predicted equivalencies.
cassava (Howe and others 2009) and carrots (Mills and others                                   vars is currently underway, especially on β-carotene stability after
2007), respectively. Bioefficacy studies in humans fed bioforti-                               processing. Carotenoid-biofortified cassava is effective in main-
fied cassava need to be done to determine the influence of cis-                                taining vitamin A status in an animal model. Continued efforts to
β-carotene content formed during processing on the vitamin A                                   improve its nutritional value are important because cassava is a
value.                                                                                         staple food for many people in developing countries.
   Measures of β-carotene absorption efficiency and conversion
to vitamin A from cassava are important parameters to study
to support the development of carotenoid-biofortified cassava.
                                                                                               Acknowledgments
Carotenoid-biofortified cassava root (yellow cassava) was as effi-                                This study is in partial fulfillment of the requirements for JAM to
cient as β-carotene supplements in maintaining vitamin A status                                obtain the Diplôme d’Ingénieur Agronome (equivalent to a mas-
in a Mongolian gerbil model (Howe and others 2009). Indeed, no                                 ter’s degree of Agronomic Engineering) from the Ecole Nationale
differences were observed for total vitamin A content in the livers                            Supérieure Agronomique of Montpellier SupAgro, France. The
of gerbils fed white cassava with daily β-carotene supplements                                 authors thank Julie Howe (Auburn Univ., Ala., U.S.A.) while at
or 45% high β-carotene cassava flour. Moreover, β-carotene was                                 the Univ. of Wisconsin-Madison for encouraging JAM during the
also found in the livers of gerbils supplemented with β-carotene                               thesis preparation stages and Harold Furr for assistance in edit-
in oil or fed 45% high β-carotene cassava, which implies that                                  ing the manuscript. We also thank Bussie Maziya-Dixon (IITA,
the gerbils had adequate vitamin A status. In addition, vitamin A-                             Ibadan, Nigeria) for providing the maps of utilization and con-
depleted gerbils fed about 15% and 30% high-β-carotene cassava                                 sumption; Bonnie McClafferty (HarvestPlus, Washington, D.C.,
flour had similar vitamin A liver content, but more β-carotene was                             U.S.A.) and JV Meenakshi (HarvestPlus, New Delhi, India) for en-
stored in the livers of the group fed 30% high-β-carotene cassava                              couragement and coordination; and Christine Hotz (HarvestPlus,
(Howe and others 2009).                                                                        Ottawa, Canada) for helpful insights. This review was sponsored
   Regarding the bioavailability of carotene from cassava leaves,                              in part by HarvestPlus contract nr 8037 and Hatch Wisconsin
Wistar rats fed synthetic β-carotene or cassava leaf powder had                                Agricultural Experiment Station WIS04975.
similar growth and tissue weight (Siqueira and others 2007).
However, β-carotene absorption was lower from cassava leaf
powder than from synthetic β-carotene. This might be due to                                   References
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