Received: 15 November 2018

| Revised: 8 February 2019 | Accepted: 10 February 2019

DOI: 10.1002/fsn3.1002

ORIGINAL RESEARCH

Nutritional potential of wild sorghum: Grain quality of
Sudanese wild sorghum genotypes (Sorghum bicolor L. Moench)
Tilal Sayed Abdelhalim1 | Nasrein Mohamed Kamal2,3 | Amro B. Hassan4
1White Nile Research Station, Agricultural
Research Corporation, Kosti, Sudan

Abstract

2

In the last decades, deficiency of macro‐ and micronutrients was considered as a seri-

Biotechnology and Biosafety ResearchCenter, Agricultural Research Corporation,
Shambat, Khartoum North, Sudan
3

Arid Land Research Center, Tottori
University, Tottori, Japan

4

Environment and Natural Resource
and Desertification Research Institute(ENDRI), National Center for Research,
Khartoum, Sudan
Correspondence
Amro B. Hassan, Environment and Natural
Resource and Desertification ResearchInstitute (ENDRI), National Center for
Research, Khartoum, Sudan.
Email: amrobabiker@yahoo.com

ous problem associated with the increase in the human population. To meet the increased demand for food consumption, the wild relative plant might serve as animportant source of new genetic material for increasing macro‐ and micronutrients.
To investigate this, the variations in protein content, in vitro protein digestibility, tannin content, phytic acid content, total polyphenol content, and total and bioavailability of minerals were studied in grains of ten wild sorghums and two released sorghum
cultivars. The results showed significant differences (p ≤ 0.05) in all quality tests
among the genotypes. The highest percentage of total protein contents and in vitro
protein digestibility were encountered in the grains of PQ‐434 (14.6%) and the released cultivar AG8 (49.8%), respectively, while the highest concentrations of total
and bioavailable iron were found in the grains of Almahkara (3.17 mg/100 g) and
Abusabiba (92.8 mg/100 g), respectively. The grains of wild sorghum genotype Adar
Umbatikh grains were found to possess higher total zinc contents. The PCA identified
only five components of eigenvalues greater than one and cumulatively accounted
for 88% of the total variation. It could be concluded that Almahkara and PQ‐434
could be used as potential sources for iron and protein sorghum biofortification, respectively. Results from this study might be used in the development of new value‐
added products from wild sorghum grains by‐products.
KEYWORDS

biofortification, genotypes, grains, micronutrients, wild sorghums

1 | I NTRO D U C TI O Nmake the crop progressively more relevant for food security in viewof climate change. In addition, it is a crop for poor communities.

Sorghum (Sorghum bicolor L. Moench, 2n = 2x = 20) is a globally im-

In Sudan, sorghum is a multipurpose crop and cultivated in al-

portant food security crop, particularly in arid and semiarid environ-most all regions by subsistence farmers for wide use. It has beenments. It is the main staple food for about 300 million people who

used to prepare different kinds of traditional food such as leavened

live in the semiarid tropics (Orr, Gierend, & Swamikannu, 2016). The

bread “kisra,” porridge “Asida,” and animal feed and to prepare local

crop is a C4 species with high photosynthetic capacity and inherent

beverages “Abraih” (Dirar, 1994). Sorghum grains are also consid-

high biomass yield potential. Its high levels of tolerance to droughtered as one of the major components of livestock and poultry feed.

and high temperature, and adaptation to inherent low soil fertility

Further, the stalk is also used as animal feed and for house and fenceThis is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium,provided the original work is properly cited.
© 2019 The Authors. Food Science & Nutrition published by Wiley Periodicals, Inc.
Food Sci Nutr. 2019;1–11.

﻿ www.foodscience-nutrition.com

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ABDELHALIM et al.

construction. The grain is characterized by its high starch, protein,micronutrients, and crude fiber but low in fat (Kumar et al., 2015). It
has been reported that majority of the Sudanese low‐income people
in rural areas, where both physical and economic access to nutrient‐rich foods are limited, depend on sorghum for their dietary energy
and micronutrient requirement (Yonos, 2001).
Micronutrient malnutrition causes blindness and anemia (evendeath) in more than half of the world's population, especially among
women of reproductive age, pregnant and lactating women, andpreschool children (Bailey, West, & Black, 2015; Graham, Welch, &Bouis, 2001). Recently, the UN Millennium Development Goal progress report for Sudan revealed that 31 percent of children under theage of five in Sudan are moderately or severely underweight (Musa,
Musa, Ali, & Mustafa, 2014). In the study of Abdelrahim et al. (2009),
anemia during pregnancy was found to be a major health problem in
Sudan, affecting around half of the women in some areas. It is a risk
factor for maternal and perinatal morbidity and mortality. Therefore,
enhancement of grain nutrients (biofortification), either agronomically (application of mineral fertilizers) or genetically (breeding), isviewed the most promising and cost‐effective approach to combat
malnutrition and related health problems (Cakmak, 2007).
Biofortification of sorghum grain through genetic strategies is a
powerful approach for changing the nutrient balance in the humandiet on a large scale. In the context of developing fortified foods
for low‐income consumers, different studies have revealed a narrow
range of genetic diversity for sorghum grain micronutrient contents,
with Fe and Zn contents (Ashok Kumar, Anuradha, & Ramaiah, 2013;
Kumar et al., 2015). The narrow genetic diversity for grain minerals
in modern cultivars could be attributed to eroded genetic diversity
by domestication and breeding processes. Therefore, efforts must
be devoted to identify valuable alleles which have been “left behind”
in the wild ancestors of crop plants and to reintroduce them into cultivated crops (Tanksley & McCouch, 1997). Interestingly, wild wheat
genotypes were found to possess high concentrations of micronutrients in grains compared to the domesticated cultivars (Monasterio& Graham, 2000). Furthermore, Peleg et al. (2008) showed large
genetic variation among 22 wild emmer wheat accessions for Zn,
Fe, and protein concentrations. The authors suggested that crop
wild relatives might serve as an important source for increasing
micronutrient concentration (Peleg et al., 2008). Hence, exploiting
the genetic variation of wild sorghum collections in Sudan for theirgrain minerals and protein contents might attract special interest forseveral reasons. Since Sudan is within the geographical range where
sorghum is postulated to be domesticated for the first time (Assaret al., 2005), where the largest genetic variation exists. To our best
knowledge, no reports on minerals and protein contents have been
published for wild and weedy sorghum. The overall objective of thisstudy was, therefore, to explore the potential of wild sorghums assources for improving the nutritional quality of grain. To do so, ten
Sudanese wild sorghum genotypes and two released cultivars werescreened in this study for antinutritional factor contents, crude andin vitro protein digestibility, and eventually total and bioavailable calcium, phosphorus, iron, and zinc contents.

2 | M ATE R I A L S A N D M E TH O DS
2.1 | Samples collection and Experimental setup
Ten genotypes of wild/ weedy species of sorghum were collected,
from the border areas between Sudan, Eritrea, and Ethiopia, a place
where sorghum is postulated to be domesticated for, the first time,
5,000 years. The wild genotypes were given names according totheir areas of the collection, classified as Sorghum bicolor var verticiliflorum and Sorghum bicolor var arundnacieum. The releasedsorghum cultivars, namely Tabat and AG8, were obtained from thesorghum breeding program at the Agricultural Research Corporation,
Wad Medani, Sudan. To affirm the homogeneity, the wild sorghum
genotypes were cultivated for three consecutive successive cycles
using a single seed descent method of plant breeding to affirm thehomogeneity. The field experiment was conducted at the White Nile
Research Farm, White Nile State, Sudan, during 2014 and 2015 cropping seasons: Latitude 13° 10ʹ N, longitude 32° 40' E, and altitude
410 meters above sea level. The climate of the locality is semiarid
with a cool dry winter and a hot summer. The rainy season extends
from July to October with peak monthly rainfall in August. The meanannual rainfall is around 330 mm. The soil of the farm is characterizedby high clay content, heavily cracking vertisols. The genotypes weresown in a two‐row plot of four meters length with the plant‐to‐plantdistance of 20 cm and row–row spacing of 80 cm. The treatments
were arranged in a randomized complete block design (RCBD) withthree replications. N fertilizer in form of urea was applied uniformly
at a rate of 43 kg N/ha once time. Panicles were harvested andthreshed from cultivars and wild genotypes at physiological maturity.
Due to the frequent shattering of spikes in various wild genotypes,
the collection of mature spikes had to be done repeatedly at different
intervals over 3–14 days. Due to tough glumes and hard threshing inwild species, the grains had to be taken out manually.

2.2 | Sample preparation
Grains of the wild sorghum and released sorghum genotypes usedin this study were carefully cleaned and freed from foreign materials. The whole grain of the samples was milled in the laboratory
miller (Retsch, Germany) using 0.4‐mm sieve to produce fine flour.
The milled samples were then packaged in a polyethylene bag andkept into a desiccator at room temperature during the analysis. All
chemicals used in this study were of analytical grade.

2.3 | Determination of crude protein and in vitro
protein digestibility
The crude protein was determined following Kjeldahl method described by AOAC (2000). The in vitro protein digestibility (IVPD) was
determined by the procedure of Maliwal (1983) cited from Monjula
and John (1991). A known weight of the sample containing 16 mg nitrogen was taken in triplicate and digested with 1 mg pepsin in 15 mlof 0.1 M HCl at 37°C for 2 hr. The reaction was stopped by addition

| 3

ABDELHALIM et al.
18
16
Protein content (%)

14
12

b

ab

b

ab

ab

b

b

2.6 | Determination of tannin content

a

ab

b

ab

ab

Tannin content of the samples was determined using the modified spectrophotometric vanillin–HCl method according to Price,

10

Socoyoc, and Butter (1978) by using 200 mg sample. A standard

8

curve was prepared and results were expressed as catechin equiva-

6

lents, that is, amount of catechin (mg/g) which gives a color intensity

4

equivalent to that given by tannins after correcting for blank.

20

2.7 | Determination of total polyphenol
Sorghum genotypes

F I G U R E 1 Protein content (%) of released sorghum cultivar andwild sorghum genotypes grains. Values are means (±SD) of triplicate
samples. Values followed by the same letter are not significantly
different (p < 0.05) as assessed by LSD. RSC1 (AG8); RSC2 (Tabat);
WSG1 (Abukarkatita); WSG2 (Abusabiba); WSG3 (Adar Abusabiba);
WSG4 (Adar Umbatikh); WSG5 (Almahkara); WSG6 (Hamadyat);
WSG7 (Hamadyat Sharateet); WSG8 (PQ434); WSG9 (Umbatikh(boda resist)); WSG10 (Zahana)of 15 ml of 10% trichloroacetic acid (TCA). The mixture was then filtered quantitatively through Whatman No. 1 filter paper. The TCA‐

Total polyphenols were determined according to Prussian Blue spectrophotometric method (Price et al., 1978). Sixty milligrams of theground sample was shaken manually for 1 min in 3.0 ml methanol. The
mixture was filtered. The filtrate was mixed with 50 ml distilled waterand analyzed within an hour. About 3.0 ml of 0.1 M FeCl3 in 0.1 M HCl
was added to 1.0 ml of the filtrate followed immediately by timed addition of 3.0 ml of freshly prepared K3Fe(CN)6. The absorbance wasmonitored on a spectrophotometer at 720 nm after 10 min from theaddition of 3.0 ml of 0.1 M FeCl3 and 3.0 ml of 0.008 M K3Fe(CN)6.

2.8 | Determination of phytic acid content

soluble fraction was assayed for nitrogen using the micro‐Kjeldahl

Determination of phytic acid content was carried out by the method

method (AOAC, 2000).

described by Wheeler and Ferrel (1971) by using 2.0 g dried sam-in vitro Protein digestibility % =digestible protein
× 100.
total protein

ple. A standard curve was prepared, and results were expressed as
Fe (NO3)3 equivalent. Phytate phosphorus was calculated from thestandard curve assuming a 4:6 iron‐to‐phosphorus molar ratio.

2.4 | Determination of total mineral
Mineral content of the grains was determined following the method of
Chapman and Pratt (1982). Each sample was burnt in a muffle furnace
at 550°C and then placed in a sand bath for 10 min after the additionof 5 ml of 5N HCl. The solution was carefully filtered in a 100‐ml volumetric flask, and finally, distilled water was added to make up to mark.
The extracts were stored in bottles for further analysis. Fe and Zn contents were determined using atomic absorption spectrophotometer.
Phosphorus was determined by spectrophotometric method using
molybdovanadate, and the absorbance was measured at 730 nm.

2.5 | Determination of minerals’ HCl extractability(in vitro bioavailability)

2.9 | Statistical analysis
Data were tested for normality of distribution and homogeneityof variances using Shapiro–Wilk and Levene's tests, respectively.
Differences between treatments were measured by ANOVA, using
STAR—Statistical Tool for Agricultural Research (version 1.0; http://bbi.irri.org/). Multiple comparisons were analyzed using Tukey range
test. Broad‐sense heritability (H2) of the mineral elements and protein was estimated from the analysis of variance following Nyquist
and Baker (1991). The formula used was H2 = VG/VP. PCA was performed using STAR software (STAR, version 1.4. International Rice
Research Institute (IRRI), Los Baños, The Philippines; http://bbi.irri.
org/products). Cluster analysis based on Euclidean distance was performed in order to employ the associations among the traits for assigning the germplasm lines and varieties to different clusters.

Minerals in the samples were extracted by the method described by
Chauhan and Mahjan (1988). About 1.0 g of the sample was shaken
with 10 ml of 0.03 M HCl for 3 hr at 37°C and then filtered. The clear
extract obtained was oven‐dried at 60°C and then acid‐digested according to Chapman and Pratt (1982). HCl extractability (%) of themineral was determined as follows:

3 | R E S U LT S A N D D I S CU S S I O N
3.1 | Crude protein content and in vitro protein
digestibility (IVPD) of the wild sorghum genotypes
and released sorghum cultivars

HCl extractability % =(mineral extractable in 0.03

Figure 1 shows the crude protein content of the sorghum genotypes

N HCl ÷ total mineral) × 100.

grains. The analysis of these genotypes showed significant (p < 0.05)

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ABDELHALIM et al.

genetic variation for the total protein content. It was varied between
10.2% and 14.6%. Among all tested sorghum genotypes, the highest
percentage of total protein content was encountered in the grains of

3.2 | Total and available mineral of the wild sorghum
genotypes and released sorghum cultivars

PQ‐434 (14.6%), whereas the lowest one was demonstrated in the

The total content and availability of Ca, P, Fe, and Zn of the wild

grains of AG8 cultivar (10.3%). The wild sorghum genotypes PQ‐434

sorghum genotypes and released sorghum cultivars are shown indisplayed significantly higher total protein contents compared to theTable 1. It was clearly observed that the total and bioavailability ofcorresponding released cultivar AG8 (Figure 1).

the elements varied among the sorghum genotypes. However, theIn relation to the in vitro protein digestibility (IVPD), it should beobtained data indicated that most of the studied genotypes grains

noted that significant (p < 0.05) variation was observed among theare rich with the major element. On the other hand, significant

wild sorghum genotypes and released cultivars. The maximum in vitro(p < 0.05) variation in the total and available content was observed

protein digestibility was determined in grains of the released culti-among the genotypes (Table 1).

var AG8 (49.8%), followed by the wild genotype Almahkara (47.4%),

Across all wild sorghum genotypes, the total Ca content of thewhereas the lowest content was obtained in grains of the wild gen-grains varied from 0.5 mg/g for Hamadyat Sharateet genotype tootype Umbatikh boda resist (29.2%). The released sorghum cultivar

2.7 mg/g for the Umbatikh (boda resist genotype), while it was foundAG8 showed significantly higher in vitro protein digestibility com-

to be 2.2 and 1.5 mg/g for the released cultivars AG8 and Tabat,pared to those of wild sorghum genotypes Abusabiba, Hamadyat,

respectively. On the other hand, the bioavailability of Ca was also

PQ‐434, Umbatikh boda resist (29.2%), and Zahana. However, thevaried significantly (p < 0.05) among the genotypes. The wild gen-wild sorghum genotypes Adar Abusabiba, Adar Umbatikh, Almahkara,otypes Zahana, Almahkara, Adar Abusabiba, and Abukarkatita con-and Hamadyat Sharateet displayed similar in vitro protein digestibility

tain higher values of Ca availability compared to those of released

compared to that of the released cultivars AG8 and Tabat (Figure 2).

sorghum cultivars AG8 and Tabat (Table 1).

From obtained results, it was clearly observed that most of theResult presented in Table 1 shows that the P content is varied

wild sorghum genotypes grains contain relatively high protein con-across the genotypes; however, no significant (p < 0.05) differences

tent with high digestibility rate. Since, protein is an essential element

were observed in P content between the genotypes. The max-of the food of animals and human and supplies the required amino

imum concentrations of total phosphorus were found in grains ofacids (Okareh, Adeolu, & Adepoju, 2015). The crude protein of thewild sorghum genotype PQ‐434 (1.98 mg/g) followed by Almahkara

wild sorghum genotypes might serve as an important source of new(1.92 mg/g), whereas the lowest concentration was obtained in

genetic material as well as fortified ingredients for enhancing thegrains of wild sorghum accession Umbatikh boda resist (1.13 mg/g).

nutritional value of grains; subsequently, most of the studied geno-

Across all sorghum genotypes, the bioavailable phosphorus con-types contain higher protein content in comparison with Sudanesetents of the grains varied from 8.5% for the wild sorghum acces-sorghum cultivars Gadambalia, Tabat, and Wad ahmed (Ahmed,

sion Umbatikh boda resist to 27.8% for the wild sorghum genotype

Abdalla, Inoue, Ping, & Babiker, 2014; Elbashir, Mustafa, Tinay, &Zahana with an average of 17.2% (Table 2). The grains of wild sor-

Babiker, 2008).

ghum genotype Zahana were found to contain significantly higher
bioavailable phosphorus contents compared to those of wild sorghum accessions Abukarkatita, Adar Umbatikh, Hamadyat, PQ‐434,

60

IVPD (%)

50
40aabcd bcd

de

abcd

abc

and Umbatikh boda resist. Nonetheless, the bioavailable phosphorus

ab
cd

abcd

bcd

bcd

were significantly higher than those of wild sorghum genotypes
e30

contents of wild sorghum genotypes Abusabiba and Adar Abusabiba
Abukarkatita and Umbatikh boda resist.
Grain mineral analysis showed a significant (p < 0.05) genetic

20

variation for total and available iron and zinc among the sorghum

10

genotypes (Table 1). The total iron concentrations of the grains var-0

ied from 1.18 to 1.91 mg/100 g with overall mean of 1.25 mg/100 g.
Across all sorghum genotypes, Umbatikh (boda resist) grains wereSorghum genotypes

F I G U R E 2 In vitro protein digestibility (%) of released sorghum
cultivar and wild sorghum genotypes grains. Values are means (±SD)
of triplicate samples. Values followed by the same letter are notsignificantly different (p < 0.05) as assessed by LSD. RSC1 (AG8);
RSC2 (Tabat); WSG1 (Abukarkatita); WSG2 (Abusabiba); WSG3
(Adar Abusabiba); WSG4 (Adar Umbatikh); WSG5 (Almahkara);
WSG6 (Hamadyat); WSG7 (Hamadyat Sharateet); WSG8 (PQ434);
WSG9 (Umbatikh (boda resist)); WSG10 (Zahana)found to possess significantly higher total iron concentration compared to the other genotypes. The highest availability values of iron
were recorded in grains of the wild genotype Abusabiba (92.8%)
followed by Zahana (78.5%). Grains of the wild genotype collectedfrom Adar Umbatikh area were found to possess the lowest available
iron concentration (49.7%). The wild sorghum genotype Abusabiba
exhibited significantly higher available iron concentrations compared to that of the released sorghum cultivar AG8. The releasedsorghum cultivars demonstrated significantly higher available iron

| 5Note. Values are means (±) SD of triplicate samples. Values without letters are not significantly different (p < 0.05) as assessed by Tukey. RSC1 (AG8); RSC2 (Tabat); WSG1 (Abukarkatita); WSG2 (Abusabiba);
WSG3 (Adar Abusabiba); WSG4 (Adar Umbatikh); WSG5 (Almahkara); WSG6 (Hamadyat); WSG7 (Hamadyat Sharateet); WSG8 (PQ‐434); WSG9 (Umbatikh (boda resist)); WSG10 (Zahana).

5.52

87.3 ± 2.2a75.0 ± 1.3ab

10.3
8.9
4.99
15.4
22.9
23.7
20.0
25.0
CV%

0.05
2.23
0.05
3.22
0.30
1.15
0.12
SE±

0.68 ± 0.05bcd
78.5 ± 4.9ab
1.20 ± 0.43e
27.8 ± 2.7a
1.51 ± 0.04a
27.8 ± 2.73a
1.2 ± 0.15cde
WSG10

69.4 ± 0.0abc
0.81 ± 0.03ab

0.63 ± 0.09cd
64.0 ± 7.7bc

75.5 ± 0.3ab
1.34 ± 0.34cde

1.91 ± 0.22a
8.5 ± 0.20 c

14.9 ± 2.3bc
1.98 ± 0.35a

1.13 ± 0.42a
8.5 ± 0.23e

14.9 ± 1.65cde
0.9 ± 0.15def

2.7 ± 0.30a
WSG9

WSG8

46.1 ± 0.0 de

75.0 ± 3.1ab

50.4 ± 1.3cde
0.7 ± 0.01abcd
70.9 ± 1.5bc
1.64 ± 0.25ab
18.5 ± 5.1abc
1.77 ± 0.20a
18.5 ± 1.12bcd
0.5 ± 0.10 f
WSG7

0.63 ± 0.04cd
68.4 ± 9.6bc
1.40 ± 0.23bcd
13.9 ± 2.2bc
1.58 ± 0.06a
13.9 ± 2.21de
1.1 ± 0.25cde
WSG6

37.2 ± 2.2e
0.87 ± 0.03a

0.72 ± 0.01abc
61.9 ± 4.8bc

49.7 ± 9.8c
1.61 ± 0.29ab

1.59 ± 0.09ab
19.5 ± 0.1abc

14.0 ± 3.8bc
1.52 ± 0.24a

1.92 ± 0.19a
23.1 ± 0.26ab

13.9 ± 2.21de
0.7 ± 0.30 ef

1.3 ± 0.30 cd
WSG5

1.49 ± 0.34
23.2 ± 4.1
1.70 ± 0.16
23.2 ± 4.06
0.7 ± 0.20
WSG3

WSG4

50.4 ± 2.5cde

58.0 ± 9.9bcd

52.0 ± 0.6

0.65 ± 0.01

bcd

0.61 ± 0.03c92.8 ± 0.1

bc1.30 ± 0.33
22.7 ± 4.3

aba1.51 ± 0.09

ef

WSG2

1.4 ± 0.20

22.7 ± 0.74

ab

9.3 ± 1.50
9.3 ± 1.47c

2.4 ± 0.50
WSG1

80.6 ± 9.6a

Available (%)

69.8 ± 8.6abc

cde

0.56 ± 0.16
71.2 ± 8.2

cde
a
ab

1.45 ± 0.18

ab1.32 ± 0.44acde

0.54 ± 0.01de
75.7 ± 1.9ab

abc
cde

1.28 ± 0.43de
17.1 ± 4.2abcca1.46 ± 0.01a
20.9 ± 5.88abceb

1.5 ± 0.15c
RSC2

0.45 ± 0.02e

Total (mg/100g)
Available (%)

61.2 ± 7.6bc
1.18 ± 0.23e

Total (mg/100g)
Available (%)

17.3 ± 2.3abc
1.66 ± 0.23a

Total (mg/g)
Available (%)

17.3 ± 2.28bcd
2.2 ± 0.40 b

Total (mg/g)
Sorghum genotypes

RSC1

Zn
Fe
P
Ca

Total and available Ca, P, Fe, and Zn of selected released and Sudanese wild sorghum genotypes
TA B L E 1

88.7 ± 0.6a

ABDELHALIM et al.

contents than those of wild sorghum genotypes Adar Umbatikh andAlmahkara (Table 1).
The concentration of Zn in grains of different wild sorghum genotypes and released sorghum cultivars is shown in Table 1. Across all
released sorghum cultivars, the wild sorghum Adar Umbatikh grains
were found to possess higher total zinc contents. Grains of thewild sorghum accessions Adar Umbatikh, Almahkara, and PQ‐434
demonstrated significantly higher total zinc contents than those of
the released sorghum cultivars. Similar to the total iron contents,
grains of the released sorghum cultivar AG8 were found to possess
the lowest total zinc concentration of 0.45 mg/100 g compared towild genotypes. Zinc available in the grains was found to be varied
from 37.2% for the wild sorghum Adar Umbatikh to 88.7% for the
released cultivar AG8 with overall mean of 65.7%. Grains of the wildsorghum accessions Abukarkatita, Almahkara, PQ‐434, Umbatikhboda resist, and Zahana displayed almost similar available zinc values compared to the corresponding released cultivars. Similar to Fe,
all studies wild genotypes are relatively containing high amount of
Zn compared to other sorghum cultivars; however, most of themshowed lower rate of Zn bioavailability.
Regarding to these findings, high concentration of Fe and Zn in
the studied wild sorghum genotypes might offer a potential source
for the biofortification through plant breeding program. Since biofortification is considered as the most suitable solution for the severe malnutrition problem particularly in the developing countries(Clemens, 2014; Stein, 2010).
Generally, our findings show that the wild sorghum genotypes
have high concentration of macro‐ and microelements. However, asignificant (p < 0.05) variation was observed among the genotypes.
These variations might be due to the genetic variability between
species and the effects of environment of the collection sites. The
genetic variability for content of the elements could be admitted asa desired trait of Sudanese wild sorghums. It has been reported thatamong the blackgram genotypes, the genetic variability showed significant factor on the variation of Zn and Fe bioavailability (Singh,
Kanaujia, Srivastava, Dixit, & Singh, 2017). Therefore, it is imperative to raise greater awareness for breeders to study the nutritional
status of diverse Sudanese wild sorghums in order to find genotypes
with elevated grain micronutrient concentrations as the initial stepfor a breeding program.

3.3 | Antinutritional factor content of the wildsorghum genotypes and released sorghum cultivars
In this study, the antinutritional factors in wild sorghum genotypes
and released cultivars were determined in the term of phytic acid,tannin, and total polyphenol content. In general, the antinutritional
factor contents of ten Sudanese wild sorghum accessions and two
released cultivars showed a significant (p < 0.05) genetic variation
for tannins, total polyphenols, and phytic acid. The grain tannin content ranged from 2.3 to 8.8 mg/g with an average of 5 mg/g. The
wild sorghum accessions PQ‐434, Abukarkatita, and Abusabiba gavesignificantly lower grain tannin contents and were not statistically

6|

ABDELHALIM et al.

Phytic aciddifferent compared to those of released sorghum cultivar AG8(Figure 3). However, contents of tannins were higher in wild sorghum
genotype Adar Umbatikh (8.8 mg/g) followed by Adar Abusabiba
1(8.6 mg/g) and were significantly different compared to that of the

Total polyphenol

released cultivars AG8 and Tabat.
Tannins can form cross‐linkages between proteins and othermacromolecules and render them unavailable for digestion
0.11

nins (Petterson, 2000). The antinutritive activity of tannin is due toits ability to make complexes with protein as well as with digestive

0.03

0.51

1

Tannin1(Griffiths, 1991). These inhibitory facets, in conjunction with an astringent taste, constitute the antinutritional characteristics of tan-

enzymes (Griffiiths 1999).

Available Zn−0.58

types collected from Zahana (6.7 mg/g). However, the wild sorghum
genotype PQ‐434 significantly contains the lower concentration of
the total polyphenol than the released sorghum cultivars AG8 and

0.12

1

−0.35
0.10

0.40

−0.52

1

Total Zn

polyphenol contents were recorded in grains of wild sorghum geno-

Tabat (Figure 4). Low concentration of total polyphenols may alsoenhance the bioavailability of trace elements, since they are able to

Available Fe

create linkage and form mineral complex which reduces its extracta0.27

0.05

0.09

−0.23

mate variations, tolerance to pests and diseases, and responses totion of nutrients have been reported (Duncan & Carrow, 1999).

−0.17

0.09

0.00

−0.01

1

to genetics, physiological/biochemical mechanisms, responses to cliagronomic management practices. Genetic variations in plant acquisi-

0.31−0.80**

1

Total Fe

−0.27

bility. (Abdelrahman et al., 2007). Such variations could be attributed

For all sorghum genotypes, grains’ phytic acid contents varied−0.10(Figure 5), while the wild sorghum genotypes Zahana, Almahkara, andAbukarkatita had significantly lower phytic acid contents comparedto the released sorghum cultivar Tabat (Figure 5). Phytic acid can

−0.01

0.14
−0.52

0.32

0. 12

sorghum genotype Hamadyat Sharateet with an average of 1.1 mg/g

−0.25

−0.13

0.09

0.03
0.40

−0.32

−0.06

1
0.36

1

Total P

0.30

Available P

from 0.2 mg/g for wild sorghum genotype Zahana to 2.4 mg/g for wildcomplex protein as well as trace elements and lower their availability (Carnovale, Lugaro, & Lombardi‐Boccia, 1988). Hence, low phytic

Values with *are significant at p < 0.05; **p < 0.01; ***p < 0.001.

−0.11
−0.22
−0.37
Phytic acid

−0.03

0.06
−0.57
−0.46
Total
polyphenol

−0.44

0.28

0.21
0.26

−0.46
0.00

0.04
0.35

−0.13

Available Zn

Tannin

−0.01

0.00
0.22

0.21
−0.52

−0.02
0.36

−0.13
Available Fe

Total Zn

0.07

0.97***
−0.30

−0.13
0.16

0.28
0.37

0.36

Available P

bioavailability rate of mineral particularly trace one.

Total Fe

0.37

1
−0.31

0.33
0.57*

0.35
0.45

0.56

Available Ca

1
0.02

0.56

IVPD

Total Ca

−0.17

1

Total Ca1

Total P

Available Caacid content in wild sorghum genotypes grains might result in high

IVPD
Crude protein−0.31

wild sorghum genotype PQ‐434 (1.3 mg/g), whereas the highest total

Crude protein

TA B L E 2

Phenotypic correlation among mineral elements and protein contents of selected released and Sudanese wild sorghum genotypes

The lowest total polyphenol contents were detected in grains of

In comparison with the antinutritional factors, tannin, total polyphenol, and phytic acid, in Sudanese sorghum cultivars Gadambalia,
Tabat, and Wad ahmed wild genotypes grains, contain less amount of
antinutrients than those of released sorghum cultivars (Ahmed et al.,2014; Idris, Hassan, Babiker, & Tinay, 2005). Moreover, the variationof the phytochemical compound among the wild sorghum genotypes
might be due to several factors particularly the growing environment.
It has been reported by Lu et al. (2015) that the environment played alarger role in variations of individual phytochemical component and antioxidant property of soft red winter wheat bran than did other factors.

3.4 | HJ‐biplot analysis
Principal component analysis (PCA) is a technique used to emphasize variation among the genotypes and to identify patterns in thedataset used for the analysis. In this study, the HJ‐biplot analysis was
conducted by comparing the principal component analysis (PCA) to| 7

ABDELHALIM et al.

strongly determine the multivariate interactions between the quality

improvement of mineral nutrients and protein is promising. High‐attributes together with the wild sorghum genotypes and released

yielding germplasm lines with improved micronutrient levels weresorghum cultivars in order to identify patterns in the dataset usedgrouped together (Tabat and Abusabiba). Selection and crossing offor the analysis table (Figure 6a,b). As illustrated in Figure 6a, thegenotypes from different clusters would help in bringing together

result of the first principal component analysis (PCA1) is explained

genes favorable for yield and quality traits so as to breed tailor‐

as 25.9% of the total variability among the genotypes, whereas themade varieties (Badigannavar, Girish, & Ganapathi, 2015; Shergo,

second principal component analysis (PCA2) is reported as 23.9%.

Labuschagne, Shargie, & van Biljon, 2013).

Hence, results of the PCA1 and PCA2 accounted for about 49.4%of the total variability representing a remarkable contribution of thetwo PC axes to the plotted components (Figure 6a). As reported by

3.5 | Phenotypic correlation among nutrients

Yan and Fregeau‐Reid (2008), the eccentricity of characters that

Correlation coefficients among mineral elements and protein areappear <90° angle is positively correlated, while the factors thatgiven in Table 2. The correlation among protein and the total andformed more than 90° angles were associated with negative corre-

8in the biplot. Accordingly, the data revealed that the wild sorghum
genotypes Almahkara, Hamadyat Sharateet, Adar Umbatikh, andAdar Abusabiba were commonly distant compared to the othergenotypes. On the other hand, the released sorghum cultivar Tabat
and wild sorghum genotypes Hamadyat and Abusabiba were mostly
stable in terms of mineral and protein contents (Figure 6a).

Total polyphenol (mg/g)

lation, and those who have a 90° angle do not show a correlation

Cluster analysis based on Euclidean distance was performed to

b

7
6
4

c

c

5
d

they had some common characters in the uptake, translocation,and the biological processes of this mineral element. Genotype
Almahkara was separated as a singleton accession from the restin this cluster (Figure 6b). This accession consisted of high crude

c

3
2

e

1

Sorghum genotypes

based on biplot coordinates into three main clusters (Figure 6b).

relatively similar mineral concentration. This indicated thatadlines and varieties to different clusters. Genotypes were grouped

Abukarkatita, Adar Umbatikh, and Hamadyat Sharateet revealed

ab

bc

0

employ associations among the traits for assigning the germplasm

Cluster indicated that genotypes Tabat, Abusabiba, Hamadyat,abb

F I G U R E 4 Total polyphenol (mg/g) of released sorghum cultivar
and wild sorghum genotypes grains. Values are means (±SD) oftriplicate samples. Values followed by the same letter are notsignificantly different (p < 0.05) as assessed by LSD. RSC1 (AG8);
RSC2 (Tabat); WSG1 (Abukarkatita); WSG2 (Abusabiba); WSG3
(Adar Abusabiba); WSG4 (Adar Umbatikh); WSG5 (Almahkara);
WSG6 (Hamadyat); WSG7 (Hamadyat Sharateet); WSG8 (PQ434);
WSG9 (Umbatikh (boda resist)); WSG10 (Zahana)protein and IVPD. The study on nutritional contents variation has
shown that identification of sorghum germplasm for breeding for

3

a

a

ab

8
bc

6
cd

d

cd

c

cd

b

bc

d

2

Phytic acid (mg/g)

Tannin (mg/g)

10

4

a

2.5

12

2

ab

ab

ab

1.5

bc

bc

1

bc

bcd
cd

bc

cd

0.5

d

0

0

Sorghum genotypes

F I G U R E 3 Tannin content (mg/g) of released sorghum cultivar
and wild sorghum genotypes grains. Values are means (±SD) oftriplicate samples. Values followed by the same letter are notsignificantly different (p < 0.05) as assessed by LSD. RSC1 (AG8);
RSC2 (Tabat); WSG1 (Abukarkatita); WSG2 (Abusabiba); WSG3
(Adar Abusabiba); WSG4 (Adar Umbatikh); WSG5 (Almahkara);
WSG6 (Hamadyat); WSG7 (Hamadyat Sharateet); WSG8 (PQ434);
WSG9 (Umbatikh (boda resist)); WSG10 (Zahana)Sorghum genotypes

F I G U R E 5 Phytic acid content (mg/g) of released sorghum
cultivar and wild sorghum genotypes grains. Values are means (±SD)
of triplicate samples. Values without letters are not significantly
different (p < 0.05) as assessed by LSD. RSC1 (AG8); RSC2 (Tabat);
WSG1 (Abukarkatita); WSG2 (Abusabiba); WSG3 (Adar Abusabiba);
WSG4 (Adar Umbatikh); WSG5 (Almahkara); WSG6 (Hamadyat);
WSG7 (Hamadyat Sharateet); WSG8 (PQ434); WSG9 (Umbatikh(boda resist)); WSG10 (Zahana)

8|

ABDELHALIM et al.

available Fe showed a negative and highly significant correlations

has implicated for the possibility of combining selection for these(R = −0.80**). Availability of Zn and Ca was significant and posi-

micronutrients in a single agronomic background. Genetic map-

tively correlated (R = 0.59*); Fe showed positive but not signifi-ping in different wheat populations confirmed QTL colocalizationcantly correlated with Zn and total protein (Table 2). Significant

conferring high protein, high Zn, and high Fe (Peleg et al., 2009).

correlation between Fe, Zn, and protein was observed in most of

Likewise, colocalization of QTLs for Zn and Fe concentrations

the studies (Wang, Yin, Tanaka, Tanaka, & Tsujimoto, 2011). Thishas been reported in rice (Garcia‐Oliveira, Tan, Fu, & Sun, 2009).

F I G U R E 6 (a) Principle component analysis (PCA1 and PCA2 = 48%) of crude protein content, in vitro protein digestibility (IVPD),
antinutritional factors, and mineral content of selected released and Sudanese wild sorghum genotypes. (b) Dendrogram showing theclustering pattern of 12 sorghum accessions from Sudan based on biplot coordinates.

| 9

ABDELHALIM et al.

TA B L E 3 Broad‐sense heritability (H2), chi‐square test (χ2), andChi probability of 14 measured variables of selected released andSudanese wild sorghum genotypes

variance especially for Fe and P (Table 3). Total phenol, total protein,Ca, and P were strongly affected by the environmental factors compared to the rest of the nutrients (Table 3). Low H2 values for some

H2

Chi (χ )probability

Chi‐square(χ2) test

traits indicated that there are no specific adaptation patterns for theTraits
Crude protein

0.73

0.00727.22

of these minerals in sorghum, any of the locations may be used for

IVPD

0.91

0.000

22.70

cultivation. Heritability estimates are limited to experimental ma-

Total Ca

0.53

0.137

2.20

Available Ca

0.80

0.0009

11.11

Total P

0.40

0.300

1.03

Available P

0.83

0.0003

13.23

Total Fe

0.97

0.000039.92

Available Fe

0.93

0.000027.47

Total Zn

0.92

0.000025.54

Available Zn

0.95

0.000032.23

Tannin0.94

0.000030.44

Total phenol

0.53

0.1301

2.29

Phytic acid

0.92

0.000023.60

2accessions studied, so when breeding for higher nutrient content

terial and setup and may differ widely in the same crop and same
trait (Garcia‐Oliveira et al., 2009). Heritability is a measure of genetic differences among individuals in a population, not simply ofwhether or not a trait is inherited (Gomez‐Becerra et al., 2010). As
decribed in Table 3, tannin, phytic acid, available Fe, Fe, available
Zn, Zn appeared to be highly variable trait with high values of heritability more than 85%.

4 | CO N C LU S I O NGenerally, obtained results revealed that there is genetic variation among genotypes. These variations might help to produce
sorghum cultivars with high nutritional quality through breeding

Some genes like grain protein content‐B1 gene (Tt‐NAM‐B1; Gpc‐

in the available gene tool. The results of our experiment showedB1), which is located on chromosome 6BS, can increase the Fe andthat the wild sorghum accession PQ‐434 exhibited fairly low tan-

Zn translocation from leaves to seeds resulting in increased Fe

nins, total polyphenols, and high crude protein contents compared

and Zn accumulation in seeds (Wang et al., 2011). Similar relation-

to the released sorghum cultivars, with a considerable potential

ship between protein, Fe, and Zn has been reported in sorghum

for improvement of the domesticated Sudanese sorghum. On the(Ng'uni, Geleta, Hofvander, Fatih, & Bryngelsson, 2012) and riceother hand, S. bicolor var arundinaceum collected from Almahkara(Garcia‐Oliveira et al., 2009). IVPD was positively and significantly

and PQ‐434 are to be considered as potential sources for iron andcorrelated with P (R = 0.61*) and the availability of P observed toprotein biofortification, respectively. These identified sources canbe totally linked with Ca availability (R = 0.96***). Highly positive

be utilized in wide hybridization for transferring useful genes fromsignificant correlation between IVPD and P revealed that concur-

alien species into cultivated sorghum. Moreover, to diversify andrently improvement of these nutrients is possible. Correlations

broaden the genetic base of cultivated sorghum, introgression

between traits are of great importance for the success of selec-of alien genes from wild sorghum needs to be persuaded vigor-

tion practiced in breeding programs. Zn content was positively

ously to make discernible yield advances in sorghum. In general,

correlated with Ca, suggesting the possibility of combining selec-high correlation value of wild sorghum genotypes to released sor-tion for both micronutrients in a single agronomic background.

ghum cultivars shows that these genotypes are potentially excel-

The association between traits and their contribution to diversity

lent materials to the sorghum breeders to produce high quality

can be validated by multivariate analysis. Breeding for high totalof sorghum cultivars. Therefore, concerted efforts are required

protein content in such a manner increases the concentration of

to screen more wild resources of sorghum for minerals and other

Mn, P, and Zn and vice versa. Waters and Pedersen (2009) also

nutritional traits.

reported positive correlation between sorghum grain protein and
Zn and P.

3.6 | Estimation of broad‐sense heritability

AC K N OW L E D G M E N T S
The authors wish to acknowledge White Nile Research Station,
Agricultural Research Corporation, ARC, Sudan, and Environment

Broad‐sense heritability (H2) of the mineral elements and protein

and Natural Resource and Desertification Research Institutewas estimated from the analysis of variance following Nyquist(ENDRI), National Center for Research, Sudan, for supporting thisand Baker (1991). Estimates of broad‐sense heritability (H2) ex-

study.

hibited medium to high % and ranged from 97% for P to 40% forFe (Table 3). Lower level of estimated broad‐sense heritability indicated the strong environmental effect and proportion of phenotypic variance attributable to environmental variance to genotype

C O N FL I C T O F I N T E R E S T
Authors declare that there is no conflict of interests.

10|

ABDELHALIM et al.

E T H I C A L A P P R OVA L
This work does not involve any human or animal experiments.

ORCID
Amro B. Hassan

https://orcid.org/0000-0002-7385-5259

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How to cite this article: Abdelhalim TS, Kamal NM, Hassan
AB. Nutritional potential of wild sorghum: Grain quality of
Sudanese wild sorghum genotypes (Sorghum bicolor L.
Moench). Food Sci Nutr. 2019;00:1–11. https://doi.
org/10.1002/fsn3.1002