South African Journal of Botany 124 (2019) 372–379

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South African Journal of Botany

journal homepage: www.elsevier.com/locate/sajb

Proximate, mineral, vitamin and anti-nutrient content of Celosia argentea

at three stages of maturity
O.D. Adegbaju, G.A. Otunola ⁎, A.J. Afolayan
Medicinal Plants and Economic Development Research Centre, Department of Botany, University of Fort Hare, Alice 5700, South Africa

a r t i c l e i n f o a b s t r a c t

Article history: The distribution of nutrients and metabolites in different organs and tissues is in a constant state of flux
Received 3 January 2019 throughout the growth and development of a plant. This study was designed to evaluate the nutritional com-
Received in revised form 3 May 2019 position of Celosia argentea at three different stages of maturity to establish the best time of harvest for opti-
Accepted 26 May 2019
mal nutritional benefits. Three growth stages: Pre-flowering (PRF), flowering (FLW) and post-flowering (PST)
Available online 13 June 2019
stages of two trials were investigated. Proximate, vitamins and anti-nutrients were performed using AOAC
Edited by L Sebastiani methods while minerals were determined using inductively coupled plasma-optical emission spectrometer
(AOAC). For both trials, the PRF had the highest ash (28.15 ± 0.10% and 26.22 ± 0.10%) and crude protein
Keywords: (19.50 ± 0.17% and 25.80 ± 0.20%) contents. However, the PST had the highest carbohydrate (28.51 ±
Proximate 0.20% and 36.16 ± 0.22%), crude fiber (33.41 ± 0.87% and 25.20 ± 0.20%) and energy (435.28 ± 27.6% and
Celosia argentea 426.08 ± 1.08%) but was the lowest in fat and moisture (8.43 ± 0.15% and 6.35 ± 0.09%) contents respec-
Mineral tively. In the mineral composition, there was no marked demarcation in the amount of Mg, Na and Fe. How-
Growth stages ever, the PRF, had the highest amount of Ca, K, P and Cu. Zinc was highest at the flowering stage of growth,
Anti-nutrients
while Vitamins A, C and E concentration decreased as the plant approached maturation. PRF stage had the
highest vitamin concentrations; alkaloid and saponin contents were highest at the PRF stage while oxalate
and phytate content was not dependent on growth stages. The study revealed that although, the nutrient com-
position of Celosia argentea is strongly influenced by maturity stages, the best age of harvest, would be deter-
mined by the deficiency to be corrected.
© 2019 SAAB. Published by Elsevier B.V. All rights reserved.

1. Introduction micronutrient deficiencies (Borokini et al., 2017). However, anti-

Increase in agricultural productivity towards reduction of hunger
has been recorded worldwide over the past two decades. However,
over 1 billion adults are overweight (Remans et al., 2011), more than
900 million people are undernourished and over 2 billion people are
afflicted by one or more micronutrient deficiencies (WHO, 2007; FAO,
2010). In a bid to produce enough calories necessary for a healthy living,
provision of adequate diversity of nutrients necessary for a healthy life is
often overlooked by agricultural and food systems. Arising from the
afore-mentioned, nutritional scientists have developed different
methods of introducing nutrients or food items to accommodate the
bigger picture of diet diversity by encouraging the consumption of fruits
and vegetables; at least one serving per day for essential nutrients and
energy (Barnes et al., 2013; Weaver and Marr, 2013).
Food plants such as leafy vegetables have played an important role
in human nutrition especially in the aspect of food security and
Abbreviations: PRF, Pre-flowering; FLW, Flowering; PST, Post-flowering.
⁎ Corresponding author.
E-mail address: gotunola@ufh.ac.za (G.A. Otunola).
nutrients substances such as phytic acid, flavonoids, tannins, saponins,
oxalates and alkaloids are present in most vegetables (Aregheore,
2012). Although most leafy vegetables contain anti-nutrients, most of
these acts as antioxidants and are responsible for their therapeutic
properties (Jimoh and Afolayan, 2009).
Celosia argentea Linn. (Amaranthaceae) is a widely cultivated vege-
table in tropical and sub-tropical Africa, North America and Asia
(Yarger, 2007). It is a leading vegetable of high economic value, which
serves as a source of living for most rural vegetable farmers during the
dry season, especially in the south western part of Nigeria where it is
known as Sokoyokoto (Akinfasoye et al., 2008). This plant has high pro-
tein and vitamin contents and is also a good source of calcium, iron, car-
bohydrates and phosphorus (Ayodele and Olajide, 2011). Apart from its
nutritional properties, various ethnopharmacological studies have re-
ported its medicinal properties which include anti-inflammatory, anti-
diarrhea, anti-urolithiatic and wound healing properties (Nidavani
et al., 2014; Varadharaj and Muniyappan, 2017). Despite its nutritional
popularity in the tropical regions of Africa and Asia, it is considered as
weed and not consumed in most parts of Southern Africa and other
parts of the globe (Council, 2006).

https://doi.org/10.1016/j.sajb.2019.05.036
0254-6299/© 2019 SAAB. Published by Elsevier B.V. All rights reserved.
 O.D. Adegbaju et al. / South African Journal of Botany 124 (2019) 372–379
Plant growth and development is highly dependent on mineral nu-
trient uptake and the concentrations of these nutrients fluctuate greatly
in space and time. This fluctuation is influenced by various factors such
as; cultural practices, stage of maturity, harvesting methods and post-
harvest handling procedures as well as the interactions among these
factors (Flyman and Afolayan, 2008; Maathus and Diatloff, 2013). Min-
eral composition of each plant organ is determined by a sequence of
events that begins with bioavailability of mineral nutrients in soils and
membrane transport in the roots to the final deposition of these nutri-
ents in to one or more cellular compartments (Grusak, 2002). Plants ac-
cumulate and redistribute macro- and micronutrients throughout their
life cycle. While some nutrients are relatively immobile in plants, some
are easily redistributed to other parts, especially during abiotic stress
conditions (e.g., nutrition depletion in the soil). Some nutrients may
be translocated from mature leaves and fruits to the younger leaves.
Also, in order to optimize reproductive potential, most plants need to
remobilize nutrients from sources in a highly orchestrated way towards
flowering. This process is spatially and temporally dependent on the
stage of the plant's life cycle. Furthermore, changes in mineral nutrient
homeostasis have been reported to influence multivariate changes in
maturity of plants (Bennett et al., 2012). Most vegetables like C. argentea
are usually harvested by uprooting or repeated cutting, which invari-
ably results to an uneven distribution of nutrients in the harvested
vegetables.
The nutritional values of C. argentea have been comprehensively ex-
plored by different researchers; however, none reported the nutrient
and antinutrients compositions at different stages of maturity, except
for the reports of Adediran et al. (2015) on the influence of leaf position
on nutritional composition of C. argentea. This study was therefore de-
signed to evaluate the impact of maturity on the nutritional composi-
tion of C. argentea at three different stages to establish the best time of
harvest for optimal nutritional benefits.
2. Materials and methods
2.1. Procurement and preparation of plant materials
Mature seeds of C. argentea were obtained from an Agro shop in
Nigeria. Seeds were raised in rectangular seedling trays measuring
65 × 100 cm2 with 200 bottom holes. They were transplanted into
72 plastic pots filled with compost soil (Khanya Nursery, Alice
Eastern Cape, South Africa) at the green house of the University
of Fort Hare, Alice, 5700 Eastern Cape, South Africa. The geograph-
ical location of the study site lies at latitude 32° 47′–19° 26′ S; lon-
gitude 26° 50′–42° 306′ E and altitude of 514.70 m above sea level.
All plants were sufficiently watered every 2 days until the experi-
ment was terminated at 12 weeks after transplanting and three
randomly selected plants were harvested per row for each harvest
period. Three harvest periods of two trials: Pre-flowering (PRF,
leaves and stem were harvested when the first flower was
sighted), flowering (FLW, harvest was done when 50% of all the
plants had flowered) and post flowering (PST, harvest was done
when the flowers were few and dropping) were investigated. The
first trial was conducted from October 2017 to January 2018
while the second trial took place from March to May 2018. A
voucher specimen (Ade/med/2017/01) at maturity stage was de-
posited at the Giffen Herbarium, University of Fort Hare. The
leaves and stem were rinsed with deionized water and gently blot-
ted with paper towel, sliced into small bits and oven-dried
(LABOTEC, South Africa) at 55 °C for 72 h until constant weight
was achieved and then ground into powder by an automated
motor blender (Polymix® PX-MFC 90D Switzerland). Physio-
chemical parameters of the soil before planting were determined
for both trials using the Inductively Coupled Plasma – Optical
Emission Spectrometer (ICP-OES) as outline by Agrilasa (2008).
373
2.2. Proximate
2.2.1. Moisture content
Moisture content was measured using air-oven following methods
of Association of Official Analytical Chemists (AOAC, 2000). A material
test chamber M720 (LABOTEC, South Africa) was used to dry an
empty weighing vessel at 105 °C for 1 h (W1) and weighed (W2). The
dry sample (5 g) was then poured in to the vessel, oven dried at 105
± 1 °C until constant weight was attained. This was then cooled in a
desiccator, after which it was weighed (W3). The percentage moisture
was calculated as:
%Moisture content ¼ W2−W3
W2−W1  100
where W1 = weight of the empty vessel. W2 = weight of the vessel
+ sample. W3 = weight of vessel + dried sample.
2.2.2. Ash content
The ash content was determined using a dry ashing method
(Agrilasa, 2007). A porcelain crucible was dried at 105 °C for 1 h, after
cooling in a desiccator, and then weighed (W1). The samples (2 g)
were placed in the previously weighed crucible and reweighed (W2).
The crucible with its content was then ashed first at 250 °C for 1 h at
550 °C for 5 h. (Furnace E-Range, E300-P4, MET-U-ED South Africa)
and allowed to cool and the weight was taken (W3). The percentage
ash was calculated as:
%Ash content ¼ W2W3
W2W1  100
where W1 = weight of a dried porcelain crucible. W2 = weight of the
crucible + sample. W3 = weight of the crucible + ashed sample.
2.2.3. Crude lipid
Crude lipid was determined using the Soxhlet extraction technique
(AOAC, 2005). The lipid content of the sample (5 g) was extracted
using 100 mL of petroleum ether. The mixture was filtered, and its
lipid content was collected in a pre-weighed (W1) clean beaker. There-
after exhaustive lipid extraction was done on the same sample with
100 mL of petroleum ether for 24 h. It was then filtered and decanted
into (W1) beaker. The lipid content was concentrated to dryness in a
steam bath and oven dried at 40–60 °C and the beaker was reweighed
(W2). Percentage of lipid was calculated as;
ð%Þ Crude lipid ¼ Weight ofW2−W1
original sample
 100
2.2.4. Crude fiber
A modification of the acid/base digestion method described by Aina
et al. (2012) was used to determine the dietary fiber. A 5 g of sample
was digested with 100 mL of 0.25 M sulfuric acid solution by boiling
under reflux for 30 min and quickly filtered. The insoluble matter was
rinsed four times with boiling water to remove the remaining acid.
This process was repeated on the residue using 100 mL of 0.31 M so-
dium hydroxide solution. The final residue was washed with water
until it was free of base. It was then oven-dried at 100 °C, cooled in a
desiccator and weighed (C1). The weighed sample was incinerated in
a muffle furnace at 550 °C for 5 h, transferred to cool in a desiccator
and weighed (C2). The percentage crude fiber was calculated as:
ð%Þ Crude fiber ¼ Weight
C2−C1
of sample
 100
2.2.5. Determination of crude protein
The total nitrogen amount in the sample was determined following
the micro Kjeldahl method (AOAC, 2005). Digestion of the sample (2 g)
was done in a Kjeldahl flask by boiling 20 mL of concentrated H2SO4 and
a Kjeldahl digestion tablet until a clear mixture was obtained. The digest
374 O.D. Adegbaju et al. / South African Journal of Botany 124 (2019) 372–379
was filtered into 250 mL volumetric flask, made up to mark with dis-
tilled water and set up for distillation. Ammonia was steam-distilled
from the digest to which 50 mL of 45% NaOH solution has been added.
The distillate (150 mL) was collected into a conical flask containing
100 mL 0.1 N HCl and methyl orange was used as an indicator. The am-
monia reacted with the acid in the receiving flask and percentage nitro-
gen (N) was estimated by back titration against 2 M NaOH. Nitrogen
calculated using the following equation.
½ðmL standard acidN of acidÞ–ðml blankN of baseÞ–ðml std:baseN of baseÞ
Weight of sample in grams
 1 : 4007
where, N = normality, percentage crude protein was obtained by mul-
tiplying the nitrogen value by a factor of 6.25.% crude protein = Nitro-
gen in sample × 6.25.
2.2.6. Total carbohydrate content
The carbohydrate content was estimated by deducting the total
crude protein, crude fiber, ash and lipid from the total dry matter as: %
Total carbohydrate = 100 – (% Moisture content + % Total Ash + %
crude fat + % crude fiber +% crude protein).
2.2.7. Energy content
Total energy of the samples was calculated by the difference method.
The atwater factors: 4, 9 and 4 kcal were employed to calculate the ca-
loric value by summing the multiplied values for crude protein, crude
lipid and carbohydrate respectively as:
Energy value(kcal/100 g) = (crude protein × 4) + (crude lipid × 9)
+ (total carbohydrate × 4).
2.3. Elemental analysis
All elemental analyses were carried out in three replications as de-
scribed by Bvenura and Afolayan, (2012) using Inductively Coupled
Plasma-Optical Emission Spectrometer (ICP-OES; Varian 710–ES series,
SMM Instruments, Cape Town, South Africa). Wavelengths, slits and
lamp current used for the determination of nine elements were
213.9 nm, 0.5 nm, 4.0 mA (zinc); 422.7 nm, 1.2 nm, 4.0 mA (calcium);
422.7 nm, 1.2 nm, 3.0 mA (copper); 589.0 nm, 0.8 nm, 3.0 mA (sodium);
248.3 m, 0.2 nm, 6.0 mA (iron); 285.2 nm, 0.1 nm, 4.0 mA (magnesium);
279.5 nm, 0.2 nm, 5 mA (manganese); 213.6 nm, 1.0 nm, 10 mA (phos-
phorus) and 766.5 nm, 0.8 nm, 4.0 mA (potassium),respectively. The re-
sults for mineral contents were expressed as mg/100 g dry
weight (DW).
2.4. Vitamins
2.4.1. Vitamin A
The retinol content in the samples was estimated using the method
described by Onyesife et al. (2014). Briefly, 1 g of sample was macerated
with 20 mL of petroleum ether. The solution was incubated for 2 h and
filtered, evaporated to dryness and 0.2 mL of chloroform-acetic anhy-
dride (1:1 v/v) added to the residue. Thereafter, 2 mL of 30% TCA-
chloroform was added to the mixture and absorbance was measured
at 620 nm using a UV-3000PC spectrophotometer. Retinol standard
was prepared in similar fashion. The concentration of vitamin A in the
sample was extrapolated from the standard curve using the equation:
Y = 0.001x + 0.0008, R2 = 0.9969.
2.4.2. Vitamin C
Ascorbic acid (Vitamin C) was quantified using the 2,6-Dichlorophe-
nol-indo-phenol, sodium salt (DPIP) titration method as described by
Adebooye (2008). The samples (1 g) were separately homogenized
with 40 mL of a buffer solution made up of 1 g/L oxalic acid and 4 g/L so-
dium acetate anhydrous. The mixture was titrated against a solution
containing 295 mg/L DPIP and 100 mg/L sodium bicarbonate. The end-
point of the titration was identified by the disappearance of the initial
blue color and results were expressed as mg 100/g dry weight.
2.4.3. Vitamin E
The vitamin E (α-tocopherol) content of the samples was deter-
mined using the method of Njoku et al. (2015). A known weight, 1 g
of the sample was macerated with 20 mL of ethanol. The solution was
filtered with Whatman No. 1 filter paper, then 1 mL of the filtrate was
added to 100 μL of 0.2% ferric chloride in ethanol and 1.0 mL of 0.5%
α,α′-dipyridyl solution. The solution was diluted with distilled water
to make the solution up to 5 mL. Absorbance was measured at 520 nm
using UV-3000PC spectrophotometer. The concentrations of the stan-
dard solutions ranged from 10 to 100 μg/mL. The concentration of vita-
min E in the samples was extrapolated from the standard curve using
the equation: Y = 0.0086x − 0.0216, R2 = 0.9985. The vitamin E con-
tent in the samples was expressed as mg/100 g DW.
2.5. Anti-nutrient composition
2.5.1. Alkaloid content
The alkaloid content was determined according to the method de-
scribed by Omoruyi et al. (2012). Five grams of sample was macerated
in 200 mL of 10% acetic acid in ethanol. The mixture was covered and
allowed to stand for 4 h. After which the mixture was filtered, and the
filtrate concentrated in a water bath to one-fourth of its original volume.
Concentrated ammonium hydroxide was added drop wise to the con-
centrated solution until precipitation (cloudy fume) was completed.
The solution was allowed to settle, washed with dilute ammonium hy-
droxide and then filtered. The residue collected with Whatman No. 1 fil-
ter paper, was dried, weighed and the alkaloid content calculated as:
Weight of precipitate
Alkaloidð%Þ ¼ Weight of original sample
 100
2.5.2. Oxalate content
The titration method described by Day and Underwood (1986) was
used to determine the oxalate content of the sample. One gram of sam-
ple was macerated with 75 mL of 3 mol/L H2SO4 in a conical flask. The
mixture was stirred with a magnetic stirrer for 1 h and filtered. The fil-
trate (25 mL) was collected and heated to 80–90 °C then maintained at
70 °C. The hot aliquot was titrated continuously with 0.05 mol/L of
KMnO4 until the end point of a light pink color which persists for 15 s.
The oxalate content was calculated by taking 1 mL of 0.05 mol/L of
KMnO4 as equivalent to 2.2 mg oxalate.
2.5.3. Phytic acid
Phytic acid was determined as described by Damilola et al. (2013).
The sample (2 g) was weighed into a flask, macerated with 100 mL of
2% HCl and allowed to stand for 3 h. The mixture was centrifuged for
10 min at 13,000 rpm, then 25 mL of the supernatant was placed in a
separate 250 mL conical flask with 5 mL of 0.3% ammonium thiocyanate
solution as indicator and 53.5 mL of distilled water was added to the
mixture. The mixture was titrated against standard iron III chloride so-
lution (0.001 95 g of iron per mL) until an end- point of slightly brown-
ish yellow color persisted for 5 min. Phytic acid was calculated as:
Phytic acid(%) = Titer value × 0.00195 × 1.19 × 100.
2.5.4. Saponin content
Saponin content was estimated using the method previously de-
scribed by Otang et al. (2012). One gram of sample was mixed with
40 mL of 20% ethanol, homogenized and allowed to stand in a water
bath for 4 h at 55 °C. The resulting mixture was filtered using a vacuum
pump. Residue was collected and re-extracted with 20 mL of 20% etha-
nol. The filtrates form the residue were combined and reduced to 40 mL
 O.D. Adegbaju et al. / South African Journal of Botany 124 (2019) 372–379
in a water bath at 90 °C; the concentrate was transferred into a 200 mL
separating funnel, 20 mL of diethyl ether was added and mixed vigor-
ously. The lower fraction was collected while the upper layer (Ether)
was discarded. The lower fraction was re- introduced into the separat-
ing funnel and 20 mL of butan-1-ol was added, mixed vigorously,
followed by 5 mL of 5% aqueous sodium chloride. The upper fraction
(butan-1-ol) was collected and evaporated to constant weight in the
oven. The saponin content in the sample was calculated using the
equation:
%saponin content ¼ Weight of fraction
 100:
Weight of sample
2.6. Statistical analysis
All data were expressed as mean ± standard deviation and were
subjected to one-way analysis of variance (ANOVA). Where data
showed significance (p b .05), Fischer's least significance difference
(LSD) test was applied using MINITAB 17 statistical package.
3. Results
3.1. Soil composition
Table 1 revealed that the soil used for planting C. argentea at the two
trials had all the elements necessary for normal growth and develop-
ment of the plant.
3.2. Proximate composition
The proximate analysis of C. argentea at different growth stages of
the two trials is represented in Table 2. The moisture content ranged
from 8.43 ± 0.15 to 10.42 ± 1.33% in the first trial and from 6.35 ±
0.09 to 9.42 ± 0.30% in the second trial. There was no significant differ-
ence in moisture content at all stages of growth in the first trial. The ash
content was highest at the PRF stage (28.20 ± 0.10%) of the first trial
and lowest at the PST (15.51 ± 0.02%) of the second trial. The fat content
ranged from 2.01 ± 0.30 to 2.50 ± 0.03% in the first trial and from 3.07
± 0.14 to 5.70 ± 0.00% in the second trial. There was no significant dif-
ference in the fat content at all the growth stages of the first trial. Crude
fiber content ranged from 22.40 ± 0.54 to 33.41 ± 0.87% in the first trail
and from 15.50 ± 0.15 to 25.20 ± 0.20% in the second trial. The percent-
age crude protein ranged from 10.61 ± 0.34 to 19.50 ± 017% in the first
trial and from 13.80 ± 0.13 to 25.80 ± 0.30% in the second trial. The car-
bohydrate content ranged from 17.85 ± 0.44 to 28.50 ± 0.20 in the first
trial and from 18.25 ± 0.50 to 36.20 ± 0.22% in the second trial. The
overall estimated energy value for both trials ranged from 315.36 ±
0.40% in the PRF stage of the second trial to 435.30 ± 1.06% of the PST
stage of the first trial. There was no significant difference in the percent-
age energy composition at all growth stages of the first trial. For both tri-
als, the PRF stage of growth had the highest ash and crude protein
content, while the PST stage had the highest carbohydrate, energy,
fiber but the lowest fat and moisture contents.
3.3. Mineral composition
Table 3 shows the result of the mineral composition of C. argentea at
different growth stage of two trials. Calcium concentration was highest
during the PRF stage and lowest at the PST stage in both trials. The FLW
stage of the first trial had the highest magnesium content of
Table 1
Mineral and physiochemical components of soil before planting C. argentea for both trials.
Samples Sample density g/mL P mg/L
First trial pre-planting soil 0.71 90
Second trial pre-planting soil 0.80 140
375
765 mg/100 g while the PRF stage of the second trial had the highest
magnesium composition of 835 mg/100 g. However, there was no sig-
nificant difference in the magnesium concentration at the three stages
of growth in both trials.
Similarly, no significant difference was observed in all the growth
phases of both trials for sodium. However, the PST stage of the first
(65 mg/100 g) and second trial (87 mg/100 g) had the highest sodium
content. Phosphorus content was highest at the PRE stage
(845 mg/100 g) of the first trial. It however decreased as it tends to-
wards maturity in the first trial. In the second trial, Table 3 shows the re-
sult of the mineral composition of C. argentea at different growth stage
of two trials. Calcium concentration was highest in the PRE stage and
lowest in the PST stage in both trials. The FLW stage of the first trial
had the highest magnesium content of 765 mg/100 g while the PRE
stage of the second trial had the highest magnesium content of
835 mg/100 g. The phosphorus content of the plant was not affected
by age as the composition (505 mg/100 g) was the same in all the
growth stages. The zinc content was highest in the FLW stage of the
first (12.30 mg/100 g) and second trial (8.7 mg/100 g). Statistically, no
significant difference was recorded in the zinc composition of C.
argentea at all growth stages of the second trial (Table 3). The highest
manganese composition of C. argentea in the first trial was recorded in
the PST stage (7.65 mg/100 g). While the highest composition for the
second trial was recorded at the PRE stage (6.15 mg/100 g). Also,
there was no statistical difference in the manganese content of the sec-
ond trial. At all growth stages, copper content was highest at the PRF
stage of growth for both trials (1.74 mg/100 g for first trial and
1.45 g/100 g for the second trial); and was not significantly different
at all growth stages of the second trial. The amount of Iron in the plant
ranged from 15.8 to 21.3 mg/100 g for the first trial and 12.5 to
13.1 mg/100 g in the second trial. Although, no significant difference
was observed in all the three growth stages in both trials, PST had the
lowest iron content, while potassium content ranged from 5995 to
10,340 mg/100 g in the first trial and from 5740 to 10,160 mg/100 g in
the second trial and the PRE stage had the highest potassium content
in both trials.
3.4. Vitamin content
Table 4 shows the vitamin contents of C. argentea at three different
growth stages of two trials. Vitamin A content of the plant samples
ranged from 4.13 mg/g DW (PST first trial) to 9.73 mg/g DW (PRE sec-
ond trial), while vitamin C ranged from 2.31 mg/g DW (PST stage of
the first and second trial) to 4.10 mg/g DW (PRE stage second trial).
There was no significant difference in the vitamin C content of the PRE
and FLW stage in both trials. Vitamin E was highest (8.68 mg/g) at the
PRE stage of growth of the second trial and lowest at the PST stage of
growth of the same trial. Overall, the PRE stage of growth had the
highest vitamin A, C and E content at both trials. NOTE: in Table 4 vita-
min A content is in (μg retinol/100 g DW), vitamin C in (mg ascorbic
acid/100 g DW) and vitamin E in (mg α-tocopherol/100 g DW), units
of these three vitamins are not in mg/g DW, therefore they cannot be
compared.
Anti-nutrient composition
Table 5 shows that phytate was the highest with a percentage value
of 8.57 ± 0.06% at the PRE stage of the first trial, followed by saponin
(4.85 ± 0.21%) at the FLW stage of the first trial and oxalate (3.70 ±
0.21%) at the PRE stage of the second trial. Alkaloid content was lowest
(0.26 ± 0.04%) at the PST stage of the second trial. For both trials, the
K mg/L Ca mg/L Mg mg/L pH (KCl) Zn mg/L Mn mg/L Cu mg/L
736 3057 783 4.92 25.7 77 5.2
1626 4272 976 6.61 27.5 3 6.2
376 O.D. Adegbaju et al. / South African Journal of Botany 124 (2019) 372–379

Table 2
Proximate composition of Celosia argentea dry weight (DW) at different growth stages of two trials.

Growth stages Carbohydrate (%) Moisture (%) Ash (%) Crude protein (%) Crude fiber (%) Crude fat (%) Energy (%)

First trial Pre-flowering 17.85 ± 0.44b 9.70 ± 0.37a 28.20 ± 0.10a 19.50 ± 0.17a 22.40 ± 0.54c 2.50 ± 0.13a 350.5 ± 0.80a
Flowering 19.80 ± 2.70b 10.42 ± 1.33a 26.25 ± 1.00b 17.24 ± 0.31b 23.83 ± 3.92b 2.50 ± 0.03a 362.68 ± 2.60a
Post-flowering 28.51 ± 0.20a 8.43 ± 0.15a 17.03 ± 0.23c 10.61 ± 0.34c 33.41 ± 0.87a 2.01 ± 0.30a 435.30 ± 1.06a
Second trial Pre-flowering 18.30 ± 0.50c 8.60 ± 0.00b 26.22 ± 0.10a 25.80 ± 0.30a 15.50 ± 0.15c 5.70 ± 0.00a 315.36 ± 0.40c
Flowering 27.01 ± 0.30b 9.42 ± 0.30a 20.27 ± 0.10b 18.50 ± 0.10b 21.18 ± 0.90b 3.65 ± 0.30b 372.58 ± 0.90b
Post-flowering 36.20 ± 0.22a 6.35 ± 0.09c 15.51 ± 0.02c 13.80 ± 0.13c 25.20 ± 0.20a 3.07 ± 0.14b 426.08 ± 1.10a

Values shown are mean ± SD; different letters along a column represent significant differences at p b .05 among the growth stages.

FLW stage had the lowest oxalate content, while the PST had the lowest
alkaloid, phytate and saponins content.
4. Discussion
The distribution of nutrients and metabolites in different organs and
tissues is in a constant state of flux throughout the growth and develop-
ment of a plant, leading to profound changes at crucial stages during
their life cycle such as vegetative, flowering and senescence and nutri-
tional content (Bennett et al., 2012). This study revealed that the nutri-
ent content of C. argentea varied across the three growth stages
investigated.
Moisture content is an index of water activity of many foods which
helps in maintaining protoplasmic content of the cell and leaf texture.
It also promotes the activity of water-soluble enzymes and co-
enzymes involved in metabolic activities of plants (Arasaretnam et al.,
2018). However, microbial contamination and chemical degradation is
associated with high moisture content in foods (Ooi et al., 2012). In all
the growth stages of both trials the moisture content of pulverized sam-
ples of C. argentea was within the acceptable range for food preserva-
tion. The relatively low moisture content observed at the PST stage of
growth of the first and second trials indicates that the plant could pos-
sess a longer shelf life at that stage of growth. This result is comparable
to the findings of Ayodele and Olajide (2011) who reported the value of
8.84% for the moisture content of the same species.
The ash content of any food sample depicts the quality of its elemen-
tal composition (Hofman et al., 2002). At the pre-flowering stage, high
levels of total ash were recorded for both trials. The ash content in this
study was higher than those reported by Ilodibia et al. (2016)
(23.91%) and Ayodele and Olajide (2011) (22.43%) for the same species.
It could be an indication that C. argentea possess more minerals at the
pre-flowering stage of growth and should be maximized for enriching
the diet with micronutrients.
Fat in food is one of the major sources of energy in human diet. It
plays a major role in the tastes perception of foods and helps the body
in the absorption of some fat-soluble vitamins. However excess fat (sat-
urated) can lead to increased cholesterol level which is a major cause of
cardiovascular disorders (Singh et al., 2009). Generally, green leafy veg-
etables are regarded as a healthy source of dietary fat because of their
low to no fat content. The very low crude fat content recorded at PST
when compared to other growth stages suggests that C. argentea
would be more suitable for the management of weight loss and some
chronic diseases associated with excess fat.
Dietary fiber plays a vital role in the regulation of bowel move-
ment, prevents coronary disease and slow down cholesterol absorp-
tion (Viuda-Martos et al., 2010). The fiber content of C. argentea at
all growth stages was higher than the values reported for Amaranthus
cruentus (8.45%) and Solanum nigrum (9.56%) (Ajayi et al., 2018). The
high fiber content recorded at the PST of C. argentea is an indication
that as plant grow older, their leaves and stem becomes woody and
more fibrous which could help in the regulation of intestinal transit,
improve dietary bulk and also decrease the risk of several metabolic
disorders caused by inadequate crude fiber intake such as colon can-
cer, obesity and diabetes in the human body (Unuofin et al., 2017
not included in references).
The protein content of most leafy vegetables ranged from 1 to 7% of
fresh weight or 8 to 30% of dry weight (Uusiku et al., 2010). From the re-
sults of this study, the protein content of C. argentea varies with the age
of the plant. The highest protein content recorded of the pre-flowering
stage of the second trial (25.80 ± 0.30%) is higher than the value re-
ported for Brassica oleracea (24.32%), A. cruentus (11.32%), S. nigrum
(15.06%) (Ajayi et al., 2018) and Moringa oleifera leaf (17.09%) (Ogbe
and Affiku, 2011). This implies that C. argentea at the pre-flowering
stage can provide more than 12% of the recommended dietary allow-
ance (RDA) of protein for children, men and women (Otten et al.,
2006). Furthermore, C. argentea protein has been reported to contain
all the major essential amino acids (Ayodele and Olajide, 2011). There-
fore, C. argentea could serve as a very good source of protein for the al-
leviation of protein malnutrition (Bauer et al., 2013). The significant
reduction of protein as the plant approaches maturation is in line with
the findings of Adediran et al. (2015). The low protein content at the
PST stage in both trials could be as a result of leaf senescence
(Table 2). A study performed on plant life cycle showed that as plant ap-
proaches senescence, photosynthesis decreases, this triggers events of
chain reactions which result in a decrease in the nitrogen contents
which is responsible for the amino acids/proteins of the plant (Avila-
Ospina et al., 2014).
Carbohydrate is the primary source of energy for bodily functions.
The highest level of total carbohydrates was recorded at PST (Table 2).
This suggests that consumption of this vegetable at the PST stage of
growth could help in providing optimal nutrition by increasing the en-
ergy content of the diet. The levels of carbohydrates in C. argentea is
higher than the value reported for A. viridis (Nisha et al., 2012) but
lower than that of Ocimum grattisimum (55.42%) (Ajayi et al., 2018).
Similarly, the overall estimated energy content of C. argentea was un-
doubtedly highest at PST because of its higher carbohydrate content.

Table 3
Elemental composition (mg/100 g) of C. argentea at three different growth stages (PRF, FLW and PST) of two trials.

Composition (mg/100 g)

Trials Growth stages Calcium Magnesium Sodium Phosphorous Potassium Zinc Manganese Copper Iron Na+/K+

First trial Pre-flowering 1540 ± 0.02a 755 ± 0.01a 45 ± 0.01a 845 ± 0.02a 10,340 ± 0.10a 11.80 ± 0.1ab 6.30 ± 0.00b 1.70 ± 0.00a 16.0 ± 0.80a 0.00 ± 0.00
Flowering 1445 ± 0.02b 765 ± 0.04a 55 ± 0.01a 770 ± 0.03a 8965 ± 0.09b 12.30 ± 0.04a 6.10 ± 0.50b 1.55 ± 0.05b 21.3 ± 1.80a 0.01 ± 0.10
Post-flowering 1365 ± 0.02c 760 ± 0.00a 65 ± 0.02a 645 ± 0.02b 5995 ± 0.05c 10.00 ± 0.70b 7.65 ± 0.50c 1.2 ± 0.10c 15.8 ± 0.00a 0.01 ± 0.11
Second trial Pre-flowering 1315 ± 0.01a 835 ± 0.01a 85 ± 0.01a 505 ± 0.01a 10,160 ± 0.11a 7.05 ± 0.50a 6.15 ± 0.05a 1.45 ± 0.05a 23.1 ± 4.20a 0.01 ± 0.00
Flowering 1160 ± 0.16a 785 ± 0.01a 85 ± 0.03a 505 ± 0.10a 9575 ± 1.43a 8.70 ± 3.60a 6.20 ± 0.80a 1.25 ± 0.30a 14.8 ± 3.00a 0.01 ± 0.01
Post-flowering 985 ± 0.01a 685 ± 0.01a 87 ± 0.01a 505 ± 0.00a 5740 ± 0.01b 4.95 ± 0.10a 4.80 ± 0.00a 0.95 ± 0.10a 12.35 ± 0.15a 0.02 ± 0.10

Values shown are mean ± SD; different letters along a column represent significant differences at p b .05 among the growth stages for each trial.
 O.D. Adegbaju et al. / South African Journal of Botany 124 (2019) 372–379
Table 4
Vitamins A, C and E contents of Celosia argentea at different growth stages of two trials.
Growth stages Vitamin A (μg retinol/100 g DW)
First trial Pre-flowering 28.64 ± 0.20b
Flowering 20.7 ± 0.10d
Post-flowering 13.72 ± 0.31e
Second trial Pre-flowering 32.42 ± 0.99a
Flowering 23.03 ± 0.40c
Post-flowering 21.05 ± 0.33d
Values shown are mean ± SD; different letters along a column represent significant differences at p b .05 among all the growth stages of both trials.
Dietary mineral elements are crucial for good and balanced human
nutrition. They support a wide variety of bodily functions, such as;
building and maintaining healthy bones and teeth, keeping the muscles
in shape and improving the functions of the heart and brain (Jéquier and
Constant, 2010). This study revealed that C. argentea at pre-flowering
stage of both trials exhibited the highest content of most minerals eval-
uated except zinc. Potassium which was the highest mineral recorded at
the pre-flowering stage, is an important mineral required in the body
for blood pressure regulation, nerve transmission, contraction, regulates
waste elimination and responsible for proper fluid balance
(Gharibzahedi and Jafari, 2017). The high potassium content at the
PRF stage of growth could have been influenced by the concentration
in the soil (Table 1), since plant growth and mineral uptake is highly de-
pendent on soil content (Aibara and Miwa, 2014). The recommended
daily allowance (RDA) of potassium for adults is 4700 mg. C. argentea
at the PRF stage is capable of contributing more than two times the
RDA for potassium This result is similar to the findings of Makobo
et al. (2010) who recorded the highest potassium content of A. cruentus
at six weeks after sowing; an age similar to the PRF stage of growth in
this present study.
The calcium content at the PRF stage of growth was also highest and
could contribute almost two times the RDA of 1000 mg/day required for
humans for calcium (Khan et al., 2011). The calcium content at all the
growth stages could also reflect the pre-planting soil which was rich
in calcium (Table 1). Calcium plays key roles in cell signaling, muscle
contraction and nerve function. It also supports the skeletal structure
by maintaining strong bones and teeth. Cells in the body use calcium
to activate certain enzymes, transport ions across the cellular mem-
branes, send and receive neurotransmitters during communication
with other cells (Sadler, 2011). Adequate intake of C. argentea could re-
duce the risk of fractures and osteoporosis in human.
Phosphorus is needed for cell growth and maintenance, helps with
kidney performance and repair. It is responsible for the formation of
adenosine triphosphate ATP and synthesis of DNA and RNA. It aids the
absorption of calcium and maintains acid–base balance (Gharibzahedi
and Jafari, 2017). In the study, the phosphorus content decreased as
the plant approached maturation; this is like the report of Makobo
et al. (2010) who recorded highest phosphorus content for A. cruentus
at four weeks old and a decrease as the plant aged. Phosphorus concen-
tration at the PRF (845 mg/100 g) endorse C. argentea as a good supple-
ment for phosphorus in the diet as 118.3 mg of C. argentea will provide
the daily 1000 mg/day amount required by adults (NHMRC, 2006).
Magnesium is required for various biochemical reactions such as the
Table 5
Anti-nutrient content of Celosia argentea at different growth stages of two trials.
Growth stages Oxalate (%)
First trial Pre-flowering 1.54 ± 0.18b
Flowering 1.25 ± 0.21d
Post-flowering 2.64 ± 0.18c
Second trial Pre-flowering 3.70 ± 0.21a
Flowering 2.42 ± 0.18c
Post-flowering 3.08 ± 0.18d
Values shown are mean ± SD; different letters along a column represent significant differences at p b .05 among all the growth stages of both trials.
377
Vitamin C (mg ascorbic acid/100 g DW) Vitamin E (mg α-tocopherol/100 g DW)
7.70 ± 0.00a 8.06 ± 0.18c
6.70 ± 0.90a 7.26 ± 0.15d
4.62 ± 0.13b 5.90 ± 0.05e
8.21 ± 0.90a 12.96 ± 0.72a
7.69 ± 1.53a 11.76 ± 0.04b
4.62 ± 1.53b 4.60 ± 0.05f
regulation of protein synthesis, muscle and nerve functions, oxidative
phosphorylation and glycolysis. It is also responsible for the formation
of protein and nerve transmission in the body (Gröber et al., 2015).
From the results obtained in this study, there was no significant differ-
ence in the magnesium content of C. argentea at all growth stages, as
such; only 53.9 mg of C. argentea will be required to meet the
450 mg/day of magnesium required for healthy living. This result cor-
roborates the findings of Adediran et al. (2015) who also reported no
significant difference in the magnesium content of C. argentea at differ-
ent age of harvest. The sodium content of the plant across developmen-
tal stages is low when compared to the 1500 mg RDA of sodium for an
adult (McCarron et al., 2013). Sodium is responsible for the mainte-
nance of electrolyte in the body. The high content of potassium in rela-
tion to the low sodium observed in this study led to a very low Na+/K+
ratio (Table 1). The ratio of Na+/K+ in any food item is a crucial factor in
nutrition, diet with Na+/K+ ratio that is less than one (Na+/K+ b 1) re-
duces the risk of hypertension (Yang et al., 2011).
Iron is one of the most common deficient micronutrients among
school children. Iron deficiency has been implicated for anemia, fatigue
and other blood related diseases (Haimi and Lerner, 2014). The iron
content of 100 g of C. argentea at the PRF stage was higher than the
RDA of 9 and 15 mg/day respectively for children and adults (Jahnen-
Dechent and Ketteler, 2012). Iron is a vital element needed for the for-
mation of hemoglobin in red blood cells. It serves as a carrier of oxygen
to the tissues from the lungs. Iron is an integrated part of some enzyme
systems in various tissues and crucial for energy production
(Gharibzahedi and Jafari, 2017). The FLW stage of growth had the
highest zinc content at both trials. This agrees with the reports of
Adediran et al. (2015) who reported zinc content of C. argentea to be
at its peak at seven weeks after sowing (like FLW stage of growth in
this study). Zinc is another important micronutrient playing important
roles in fetal development, normal growth and sexual maturation of
human beings. Deficiency in zinc could lead to low immune system
(Roohani et al., 2013). In this study, zinc content progressively declined
with advancing maturity: a trend also reported by Flyman and Afolayan
(2008) for Mormodica. balsamina and Vigna unguiculata. With the zinc
content recorded at the FLW stage, 32.52 mg and 113.82 mg of C.
argentea respectively can provide the 4 and 14 mg/day requirement of
zinc for children and adult (NHMRC, 2006).
Copper and manganese are essential trace elements that are
needed only in minute amounts by the human body for biochemical
functions. Manganese acts as a co-factor to many enzymes, it is impor-
tant for the normal functioning of the brain and proper activity of
Phytate (%) Alkaloid (%) Saponins (%)
8.57 ± 0.06a 3.18 ± 0.21b 3.10 ± 0.10c
5.71 ± 0.50c 2.67 ± 0.24d 4.85 ± 0.21a
5.69 ± 0.41c 1.98 ± 0.10e 2.82 ± 0.15c
4.04 ± 0.14d 3.34 ± 0.37a 1.82 ± 0.15d
6.99 ± 0.8b 2.85 ± 0.21c 3.91 ± 0.13b
2.73 ± 0.30e 0.26 ± 0.04f 1.00 ± 0.00e
378 O.D. Adegbaju et al. / South African Journal of Botany 124 (2019) 372–379
nervous system throughout the body (Au et al., 2008). Copper act as a
co-factor and constituents of several enzymes; it is essential for the
proper functioning of organs and stimulates the immune system to
fight infections (DiNicolantonio et al., 2018). Deficiency in copper
leads to cardiac abnormalities, while manganese deficiency results in
skeletal abnormalities, impaired growth, and abnormal lipid metabo-
lism (Au et al., 2008; DiNicolantonio et al., 2018). Copper content at
all growth stages of C. argentea could provide the RDA of 0.7, or
1.1 mg/day required for children and adults respectively (Johnson
et al., 2009 not included in references). While manganese content at
the PRF stage of growth (first trial) could contribute two times the
RDA value (NHMRC, 2006).
Vitamins are organic compounds that help the body to grow
and function properly by boosting the immune system. In this
study, the PRF of both trials had the highest amount of all the vita-
mins evaluated. Vitamin A is required for the normal functioning
of the visual system, growth and development, maintenance of ep-
ithelial cellular integrity, immune function and reproduction,
while vitamin C helps in the biosynthesis of carnitine, hormones,
collagen and iron absorption. It is a water- soluble antioxidant
which acts as a free radical scavenger and stimulates the immune
system; thus, promoting health in human beings. Severe deficiency
of vitamin A can lead to xerophthalmia while that of vitamin C can
lead to scurvy (Sommer and WHO, 1995; Locato et al., 2013). The
key biological function of vitamin E is the protection of the polyun-
saturated fatty acids of cell membranes from free-radical damage
in oxidative stress (Korchazhkina et al., 2006). The vitamin content
of C. argentea in this study decreased as the plant approached ma-
turity. A similar trend was reported by Biesiada et al. (2007) for
Leek cv. Kilima, Delikates biala and Zucchini cv Astra. This implies
that the best time to harvest C. argentea for optimum vitamin con-
tent is at the PRF stage of growth.
Antinutrients in foods interfere with the absorption of some min-
erals and other micronutrients in the digestive system which may
have a negative impact on the functioning of certain organs (Gemede
and Ratta, 2014). In this study, wide variation was observed in the con-
tent of the four antinutritional compounds evaluated at the three
growth stages.
Alkaloids are naturally occurring chemical compounds with basic
nitrogen atoms; they are active components of medicinal plants. How-
ever, high percentage of alkaloid is toxic to human and animals
(Matsuura and Fett-Neto, 2017). The highest alkaloid content at the
PRF is within the range of alkaloids content that is perceived to have
more pharmacological effect rather than toxicity (Makkar et al.,
2007). Oxalate is present in the cell sap of many of the green leafy
vegetable. It affects the human body by forming a strong chelate
with dietary calcium and other minerals thereby rendering such nutri-
ents unavailable for absorption and assimilation (Jiru and Urga, 1995).
This insoluble calcium oxalate in the crystal form is stored in the kid-
ney causing serious health-related problems called kidney stone
(Natesh et al., 2017). The oxalate content in all the growth stages
did not follow a trend in the growth stages. However, the highest
values for oxalate content of C. argentea at all the growth stages in
both trials are lower than that of A. cruentus (5.08 mg/g) and S. nigrum
(5.88 mg/g) (Ajayi et al., 2018).
Phytate is usually present as salts in grains and vegetables. Nega-
tively charged phosphate group in phytic acid is known to inhibit the ac-
tion of gastrointestinal tyrosinase, trypsin, pepsin, lipase, amylase and
essential minerals (Hendricks and Bailey 1989 not included in refer-
ences). The result of the phytate content of C. argentea in this study
did not follow a definite trend in both trials as observed with the oxalate
content. The implication of this is that the phytate content of this plant is
not affected by maturity. Phytic acid at a level of 0.035% is reported to
have a protective measure against fatty liver resulting from elevated he-
patic lipogenesis (Onomi et al., 2004). The highest phytate content (8.57
± 0.06%) is within the safe limit, since the inhibition of mineral
absorption by phytate only occurs at levels greater than 10% in a diet
(Onomi et al., 2004).
Saponins at high concentrations can affect nutrient absorption by
inhibition of metabolic and digestive enzymes as well as binding
with nutrients such as zinc (Shahidi, 1997). The saponin content re-
corded at the three growth stages in this study was low compared
to the value recorded for Vernonia amygdalina(5.20 mg/100 g) (Ladi
et al., 2016 not included in references). When saponin in a diet is
less than 10%, it is believed to be harmless to the body (Hortwitz,
2003 not included in references). The highest content of saponin re-
corded at the flowering stage of growth of both trials was within
the safe limit.
5. Conclusion
The study confirmed that maturity stages have a significant influ-
ence on the nutritional composition of C. argentea. The antinutrient con-
tents were low when compared to some other green leafy vegetables;
therefore, interference with nutrients absorption will be negligible.
This study present C. argentea as a vegetable with high potential to com-
bat the menace of nutrient deficiency, its cultivation is therefore, highly
recommended to address food and nutrient insecurity. Although, the
potential of this plant as an important source of nutritional and mineral
compounds is possible at all the three growth stages, the best stage of
harvest would be determined by the specific nutrient need.
6. Significant statement
This study provides insights into the nutritional composition of C.
argentea at three growth stages. The results reveal that maturity stages
significantly influence the nutrient content of this plant and serves as a
baseline data for the formulation of guidelines towards the best age of
harvest suitable for optimum nutrition. The consumption of C. argentea
as a green leafy vegetable and source of dietary vitamin and mineral at
all stages of growth is greatly encouraged. The plant can also be utilized
to boost the immune system. The findings of this study are relevant in
situations where a food-based strategy is sought to address specific nu-
trient and mineral deficiencies.
Author's contributions
Anthony Afolayan and Gloria Otunola conceptualized the topic;
Gloria Otunola and Oluwafunmilayo Adegbaju designed the format of
the experiment. Oluwafunmilayo Adegbaju carried out the experiment
and wrote the manuscript. Anthony Afolayan and Gloria Otunola revise
the manuscript. All the authors read the final manuscript.
Acknowledgement and funding
Authors acknowledge the financial support of Govan Mbeki
Research Development Centre, University of Fort Hare. Grant number:
C127.
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