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Amaranth: A New Millennium Crop of Nutraceutical
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Anu Rast ogi
a
& Sudhir Shukla
a
a
Nat ional Bot anical Research Inst it ut e, Lucknow, India
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Critical Reviews in Food Science and Nutrition, 53:109–125 (2013)
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ISSN: 1040-8398 / 1549-7852 online
DOI: 10.1080/10408398.2010.517876
Amaranth: A New Millennium Crop of
Nutraceutical Values
ANU RASTOGI and SUDHIR SHUKLA
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National Botanical Research Institute, Lucknow, India
The major staple food crops production is not able to fulfill food requirement of the global population due to relatively higher
population growth rate in developing countries. The research on these crops for exploring their ultimate yield potential is
currently at a plateau level. To replace the existing pressure on these major crops there is an urgent need to explore other
alternative crops having the potential to replace and fulfill the available food demand. FAO statistics reveal that there is a
high frequency of low birth weight children in the developing countries, which is primarily due to deficiency of micronutrients
in the mother’s diet. Amaranth, an underutilized crop and a cheap source of proteins, minerals, vitamin A and C, seems to be
a future crop which can substantiate this demand due to its tremendous yield potential and nutritional qualities, also recently
gained worldwide attention. Recently, current interest in amaranth also resides in the fact that it has a great amount of
genetic diversity, phenotypic plasticity, and is extremely adaptable to adverse growing conditions, resists heat and drought,
has no major disease problem, and is among the easiest of plants to grow in agriculturally marginal lands. The present
review is an effort to gather the available knowledge on various diversified fields of sciences for the future exploitation of
the crop.
Keywords Amaranthus, nutritive value, medicinal aspect, molecular, breeding
INTRODUCTION
The family Amaranthaceae is generally considered as the
“Amaranth family.” The word Amaranthus is basically derived
from the Greek word “Anthos” (Flower) which means everlasting or unwilting. At the present time it is also called a third
millennium crop plant. Based on taxonomical studies, the family
is divided into two sections (Allen, 1961), namely Amaranthus
saucer and Blitopsis dumort, with nearly an equal number of
species (Thellung, 1914). The section Amaranthus is dibasic
with x = 16 and 17 and Blitopsis with x = 17 except polyploidy, that is, A. dubius (Pal, 1972; Pal and Khoshoo, 1965;
Madhusoodanan and Pal, 1981). The genus amaranth is mainly
comprised of about 400 species among which few of them are
found worldwide. The division of species is based on their utilization method into grain amaranth, vegetable amaranth, ornamental, and weedy amaranth (Sauer, 1967). Grain amaranth has
four species, that is, A. hypocondricus, A. cruentus, A caudatus, and A. edulis associated with three weedy species A. hybridus, A. powellii, and A. quitensis (Pal and Khoshoo, 1974).
The vegetable Amaranthus belongs to the section blitopsis (Pal
and Khoshoo, 1972; Madhusoodanan and Pal, 1981). Vegetable
Address correspondence to Sudhir Shukla, 1 National Botanical Research
Institute, Lucknow, India. E-mail: s shukla31@rediffmail.com
amaranth has two major species, A. tricolor and A. lividis. The
ornamental species includes A. tricolor (Table 1). Amaranth is
a fast growing crop and because of its low production cost, it is
one of the cheapest dark green vegetable in the tropical market
and is often described as the poor man’s vegetable. Unlike the
other green vegetables, it is cultivated during summer when no
other green vegetables are available in the market (Singh and
Whitehead, 1996). The amaranth can grow under varied soil
and agroclimatic conditions (Katiyar et al., 2000; Shukla and
Singh, 2000), and is also resistant to heat and drought with no
major disease problems (Robert et al, 2008; Barrio and Anon,
2010). Besides its adaptable nature in various climatic conditions, the amaranth plant also has important nutritional and
medicinal properties (Lakshmi and Vimala, 2000). Prakash and
Pal (1991); Shukla et al. (2003; 2006) and Prakash et al. (1995)
emphasized the use of amaranth as a vegetable and grain crop
which can be a cheap alternative rich source of protein and
nutrient for poor people in developing countries.
HISTORY
The historical evidence depicted that Amaranthus domestication and cultivation came into use about 8000 years ago in
the Mayan civilization of south and Central America. The most
109
110
Table 1
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S.No
1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
11.
12.
13.
14.
15.
16.
17.
18.
19.
20.
21.
22.
23.
24.
25.
26.
27.
28.
29.
30.
31.
32.
33.
34.
35.
36.
37.
38.
39.
40.
41.
42.
43.
44.
45.
46.
47.
48.
49.
50.
51.
52.
A. RASTOGI AND S. SHUKLA
List of various species with their local name of Genus Amaranth (cited from Wikipedia)
Name of Species
Amaranthus gangeticus
Amaranthus floridanus
Amaranthus graecizans
Amaranthus fimbriatus
Amaranthus dubius
Amaranthus deflexusAmaranthus cruentus
Amaranthus crispus
Amaranthus crassipes
Amaranthus chlorostachys
Amaranthus chihuahuensis
Amaranthus caudatus
Amaranthus cannabinus
Amaranthus californicus
Amaranthus brownii
Amaranthus blitum
Amaranthus blitoides
Amaranthus bigelovii
Amaranthus australis
Amaranthus acanthochiton
Amaranthus acutilobius
Amaranthus albus
Amaranthus arenicola
Amaranthus torreyi
Amaranthus thunbergii
Amaranthus standleyanus
Amaranthus spinosus
Amaranthus scleropoides
Amaranthus rudis
Amaranthus retroflexus
Amaranthus quitensis
Amaranthus pumilus
Amaranthus pringlei
Amaranthus viridis
Amaranthus watsonii
Amaranthus wrightii
Amaranthus tuberculatus
Amaranthus tricolor
Amaranthus lineatus
Amaranthus leucocarpus
Amaranthus hypochondriacus
Amaranthus hybridus
Amaranthus greggii
Amaranthus powelii
Amaranthus polygonoides
Amaranthus paniculus
Amaranthus palmeri
Amaranthus obcordatus
Amaranthus muricatus
Amaranthus minimus
Amaranthus mantegazzianus
Amaranthus lividus
significant historical evidence supports that the amaranths was a
staple food, called huahtli grown in Mexico during the Aztec civilization (Sauer, 1950a; 1950b; Pal and Khoshoo, 1972; Early,
1977; Haughton, 1978; Lehman, 1994). The Aztecs believed
that it had magical properties which gave it strength. Due to this
belief Amaranthus was used as a grain in religious practices, and
was roughly equal to corn. But in the 1500s, the Spanish con-
Vernacular name
elephant head amaranth
Florida amaranth
fringed amaranth, fringed pigweed
spleen amaranth, khada sag
large-fruit amaranth
purple amaranth, red amaranth, Mexican grain amaranth
crispleaf amaranth
spreading amaranth
chihuahuan amaranth
love lies-bleeding, pendant amaranth, tassel flower, quilete
tidal-marsh amaranth
California amaranth, California pigweed
Brown’s amaranth
purple amaranth
mat amaranth, prostrate amaranth, prostrate pigweed
Bigelow’s amaranth
southern amaranth
greenstripe
sharp-lobe amaranth
white pigweed, prostrate pigweed, pigweed amaranth
sandhill amaranth
Torrey’s amaranth
Thunberg’s amaranth
spiny amaranth, prickly amaranth, thorny amaranth
bone-bract amaranth
tall amaranth, common water hemp
red-root amaranth, redroot pigweed, common amaranth
ataco, sangorache
seaside amaranth
Pringle’s amaranth
slender amaranth, green amaranth
Watson’s amaranth
Wright’s amaranth
rough-fruit amaranth, tall waterhemp
Joseph’s-coat
Australian amaranth
Prince-of-Wales-feather, princess feather
smooth amaranth, smooth pigweed, red amaranth
Gregg’s amaranth
green amaranth, Powell amaranth, Powell pigweed
tropical amaranth
Reuzen amarant
Palmer’s amaranth, palmer pigweed, careless weed
Trans-Pecos amaranth
African amaranth
Quinoa de Castilla
-
quistadors prohibited the growing of Amaranthus to suppress
the Aztec culture and religion which promoted its adoption and
production in the other parts of world. In the present time only
a limited amount of grain is being grown in Aztec and used
in the production of “alegria” candy after mixing with honey
(Early, 1977) and for preparing a tole, a drink (Oke, 1983). Evidence showed that the pale seeded amaranth was also grown in
111
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NUTRITIVE VALUE OF AMARANTH
Germany in the sixteenth century. By the 1700s it had spread
throughout Europe and was being used as a herb and ornamental. In the 1800s it was reportedly being grown in the mountain
valley of Nepal and parts of east Africa, and later in the nineteenth century it spread to the Himalayas and interior China
and eastern Siberia (Sauer, 1977). In Europe it was planted for
ornamental purposes and for in Africa for vegetable purposes.
Only in the Himalayan region was it eaten and maintained as a
minor cereal. During the twentieth century, it had been grown in
China, India, Africa, and Europe, as well as in North and South
America. Although the U.S. has been the leading producer of
grain amaranth which is being used in retail food products, but
the largest production area in the last decade is believed to have
been in China. The Chinese use amaranth as forage to feed hogs,
rather than harvesting it as grain.
In ancient Greece, the Amaranthus was named Chrusanthemon and Elichusos and was also sacred to Ephesian Artemis.
They supposed that amaranth had healing property and as a symbol of immorality was used in decoration of gods and tombs.
In Greek mythology, Amaranthus (a form of Amarantus) was a
hunter and king of Euboea island for which it was the eponymous hero. There was a famous temple of Artemis (Amarynthia
or Amarysia). It was believed that he was loved by the goddess
Artemis and joined her in the hunt. But he insulted Poseidon
without any reason due to which the Gods sent a giant wave
to wash him into the sea which drowned him. Artemis later
turned him into an amaranth-flower, her sacred plant. It was
also reported that Pagan tribals used Amaranthus in their burial
ceremonies.
ORIGIN AND DISTRIBUTION
About 400 species of Amaranthus are distributed throughout
the world in temperate, subtropical, and tropical climate zones
(Suma et al., 2002). A few of them are distributed worldwide
(Anon., 1992). About 20 species are found cultivated/wild in
India. Among grain types some species are considered as native
to south and Central America (Grubben and Van Stolen, 1981)
while some other types are native to Europe, Asia, Africa, and
Australia (Becker et al., 1981; Teutonico and Knorr, 1985). In
its own country of origin, grain Amaranthus (A. caudatus) is
known by various names (Table 2). India has been considered
as one of the centers of distribution of amaranth, the other center
being tropical America. Dioceous forms are generally absent in
India and the old world and found only in the new world.
There is no authenticate/concrete evidence of the origin of
Amaranthus. According to one view grain amaranth is being
cultivated in the old world from time immemorial and probably
originated there. The other view stated that it was introduced in
India (Malabar Coast) from the new world (Brazil) by early Portuguese traders around 1500 A.D. It is probable that they were
independently domesticated by different prehistoric people of
North and South America (Anon., 1992). The ornamental type
(A. tricolor) believed that it originated in India and then introduced to the new world. From species A. tricolor, several domes-
Table 2
Name of Amaranthus in other languages
S.No.
Name of Countries
1.
China
2.
3.
4.
5.
6.
7.
8.
9.
10.
11.
Japan
India
Indonesia
Malaysia
Laos
Philippines
Sri Lanka
Thailand
Vietnam
Peru
12.
13.
Bolivia
Eucador
Local Name of Amaranthus
een choy, yin choy, in-tsai, hsien tsai,
xian cai
hiyuna
Chaulai
bayam
bayam
pak hom
kulitis
thampala
pak khom hat, pak khom suan
yan yang
anchita, achos, achis,incajataco, coimi
and Kiwicha
coimi and millmi
Sangoracha, alaco
tic varieties of ornamental and vegetable types have been developed (An., 1992). The main vegetable type (A. tricolor) seems
to have originated in south Asia (Grubben and Van Stolen, 1981)
and then spread throughout the tropics and temperate areas
(Martin and Telek, 1979). A. tricolor has been extensively cultivated, primarily in southern China (Martin and Ruberte, 1979).
ARCHEOLOGY
The earliest archaeological record of this pale seeded grain
stated that it came into usage as a grain about 4000 years ago
in Tehuacan Puebla, Mexico (Pal and Khoshoo, 1974; Sauer,
1979). The oldest seeds found are of a species A. cruentus.
Some earliest seeds of A. hypocondriacus and A. cruentus are
also found from the old deposits both in caves at Tehuacan and
Mexico. Sauer (1979) observed that seeds could be domesticated
far away and earlier than those findings indicate. The seeds of
A. hypocondriacus are also identified in the cache located in a
rock shelter in the Ozark mountains of North West Arkhansas.
Archaeologists have also recorded the evidence of Amaranthus
domestication from numerous prehistoric culture sites including
Maverick County and the Hinojosa Site near Alice in Jim Wells
County, etc. The similarity between Chenopod and amaranth
seeds creates confusion for the archaeologist for its clear identification. To solve this problem the seeds are being preserved
to make positive identification up to the genus level.
BOTANICAL DISCRIPTION AND AGRONOMICAL
PRACTICES
The genus Amaranthus generally includes monoecious annuals except some dioceous form (much restricted in distribution
and has a branched and bushy appearance). The plant height
varies from 0.3 m to 5 m among the various species. Leaves are
oblong to elliptical in shape with the color ranging from light
to dark green with some expressing red pigment throughout
the genus. The inflorescence is very prominent, colorful, and
112
A. RASTOGI AND S. SHUKLA
Table 3 Brief description of habit and plant morphology with basic requirements for its agronomical practices. (Palado and Chang, 2003; Stallknert and
Schulz-Schaelfer, 1993)
Plant Description
Life form
Habit
Life span
Herb
Upright
Annual
Category
Branching
Plant attributes
Vegetables and Grain
Multi branched
Grown on small scale
Ecology
Crop cycle
Temperature requirement
Through out year
25◦ C–40◦ C
Soil depth
Soil texture
50 tonnes of wet rotten FYM per ha as basal dose before planting. After
preparing trenches, apply N:PzOs:K2O @ 50:50:50 kglha. Another
50 kg of N can be applied at regular intervals as topdressing. Spraying
1% urea immediately after each harvest
Soil fertility
shallow (4mm)
Medium, light heavy and
organic
Nutrient rich soil, with potash
and nitrogen
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Manuring
Soil drainage
Soil PH
Spacing
Climatic zone
Cultivation
Abiotic tolerance
Susceptibility (Biotic)
Plant protection
Useful parts
Temperate, tropical and sub tropical
Tolerant to high aluminium soils and arid conditions or drought tolerant
and poor fertility
damping-off disease, root rot, and caterpillars, stem borers, and leaf
webber attack
Avoid use of insecticides or fungicides.
Whole plant as leafy green vegetables and seed used directly or as flour
Light intensity
Elevation
Well dry spells
5.0–8.0
15–30 cm in rainy season
planting on raised beds
very bright and clear skies
Sea level to 3500 m
Seed rate (Sowing)
1.5–3.0 Kg/ha
Susceptibility
(Abiotic)
thrives in 30–35◦ C and
stressful conditions
terminal and contains one male flower per glomerule (of nearly
100–250 flowers) in section amaranth while small, generally
auxiliary with 10–25% male flowers per glomerule in section
Blitopsis. In the dioceous form at least two propagules (male
and female) require to disperse together (Khoshoo, NBRI silver
Jubilee). The monoceious habit with predominant outcrossing in
grain amaranth helped in their domestication (Pal and Khoshoo,
1974). The pollen grains are spherical in shape with poly ontoporate or golf ball like aperture in monoecious form. The pollen
grains of dioceous form are poly aperturate in their visible surface. There is no difference between the diameters of aperture of
both forms (Franseen et al., 2001). Seeds are small and lenticular
averaging 1–1.5 mm in diameter with 1000 seeds weight ranged
from 0.6–1.2 g (Jain and Hauptli, 1980; Saunders and Becker,
1984). The color of seeds varies species to species from pale
ivory to black (Pal and Khoshoo, 1974; Saunders and Becker,
1984; Irving et al., 1981; NAS, 1975). Seed embryo remain surrounded with the nutritive layer, thin wall perisperm cells being
full of starch grains, and the protein bodies are embedded in
lipid matrix (Coimbra et al., 1994). The prescribed agronomical
practices for better cultivation are presented and shown in Table
3 (Palado and Chang, 2003; Stallknert and Schulz-Schaelfer,
1993).
et al., 1981; Saunders and Becker, 1984). The shape of starch
granules varies species to species as small, 1–3 µ in diameter,
angular, and polygonal in shape in A. hypocondriacus (Lorenz,
1981; Saunders and Becker, 1984; Stone and Lorenz, 1984),
spherical as well as angular and polygonal with smooth surface
and size varies from 0.75 to 1.5 nm in A. cruentus (McMasters
et al., 1955; Stone and Lorenz, 1984; Qian and Kuhn, 1999;
Hoover et al., 1998), and spherical in small amount with large
irregular starch chunks in A. retroflexus (Goering, 1967). A.
hypocondriacus seeds have nonglutinous and glutinous type
starch with nearly 100% typical amylopectin (Tomita et al.,
1981; Okumo and Sakaguchi, 1981), nonglutinous type in A.
caudatus (Okumo and Sakaguchi, 1981; 1982), and glutinous
types in A. cruentus (McMasters et al., 1955). As compared
to the starch in corn and wheat, the starch of A. cruentus and
A. hypocondriacus has higher swelling power or absorbance
capacity (Resio et al., 1999), lower solubility, greater uptake,
lower susceptibility to amylases, and lower amylase content
(4.7 to 12.5%) (Qian and Kuhn, 1999; Stone and Lorenz, 1984;
Kong et al., 2009) (Table 4).
CHEMICAL COMPOSITION
Components
Starch
Starch is the main component of amaranth grain and used in
food preparations (Wu and Corke, 1999). Starch is reported to
be 48% in A. cruentus and 62% in A. hypocondriacus (Becker
Table 4 Comparison of grain amaranth with other major cereals (% dry
basis) (Pedersen et al., 1987)
Ash
Protein (N × 6.25)
Fat
Sugar
Starch
Dietary Fiber
Maize
Wheat
Sorghum
Amaranth
1.2
10.0
5.2
3.2
72.8
9.3
1.7
13.2
2.7
4.2
65.7
12.1
1.7
12.6
4.0
2.0
70.1
8.5
2.5
14.5
10.2
3.1
62.7
8.8
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NUTRITIVE VALUE OF AMARANTH
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Fat/Oil
Amaranthus oil is extracted from the seeds of two species,
that is, A. cruentus and A. hypocondriacus, which range between 4.8–8.1% (Gamel et al., 2007; He and Corke, 2003). The
hyper oxide stability test showed oil is more soluble than sunflower oil (Gamel et al., 2007). The melting point of amaranth
oil is −27◦ C. The oil is medium to light in color, clear and
pourable, at low temperature, and highly unsaturated with a delicate agreeable aroma and taste. It contains mainly non-polar
lipid compounds especially triglycerides (80.3–82.3%) with a
degree of unsaturation and a very low amount (about 9.1–10.2%)
of phospholipids (Gamel et al., 2007). Oil is also an excellent
source of omega series of fatty acids. The digestibility of oil
corresponds approximately to that of cotton but A. cruentus oil
has low digestibility. The oil also has tocols (1465.15 mg/kg)
and squalene (upto 6.8%) as compared to wheat germ oil which
has only 0.1–1.7%. Squalene, an oxidation resistant lubricant,
is unsaturated open chain fatty acid generally used in the cosmetic and skin care products production industry. A. cruentus
has 4.0–10.0% squalene (Gamel et al., 2007; Grazdiene, 2007),
A. hypocondriacus has 6.1%, A. tricolor has 5.1% while A.
edulis has 6.7% (Becker et al., 1981; He and Corke, 2003;
He et al., 2002). The oil is composed of myristic acid (upto
0.6%), palmitic acid (upto 18.7%), stearic acid (upto 5.3%),
anarchidic acid (upto 1.9%), behenic acid (upto 2.6%), oleic
acid (upto 30.5%), linolenic acid (upto 62%), and linoleic acid
(upto 40%) (Badami and Patil, 1976; Jahaniaval et al., 2000;
He and Corke, 2003; He et al., 2002). It was noticed that there
was a positive correlation between squalene yield and oil content and negative between linoleic and either of the two fatty
acids, palmitic acid and oleic acid. The ratio of saturated and
unsaturated fatty acids decreased with the maturity (He and
Corke, 2003). Amaranth oil can raise HDL cholesterol and significantly reduce non-HDL cholesterol which lowers the low
density lipoprotein cholesterol by 21–50% (Chaturvedi et al.,
2007).
The seeds of the spp. A. tricolor yield upto 4.3% oil
with 22.2% saturated and 77.8% unsaturated fatty acids
(Chidambaram and Iyer, 1941). Among the saturated acids, the
content of myristric acid remain upto 24.3%, palmitic acid upto
38.4%, stearic acid upto 32.98%, anarchidic acid upto 2.32%,
while in unsaturated acid linoleic acid upto 55.13% and oleic
acid upto 44.87%. A growth inhibitory steroid amasterol is reported in the roots which was identified as 24 methylene 20
hydroxy cholesterol-5, 7-en, 3 β-ol (Roy et al., 1982).
Table 5
Fat, the second important content present in grain amaranth,
is higher than cereal. The fat is characterized by a high content of unsaturated fatty acids and major saturated fatty acids.
Linoleic acid is a major fatty acid in oil which is more than
50% in the seeds of grain amaranth. The next higher one is oleic
acid with more than 20% followed by palmitic acid which is
about 20%. The major unsaturated fatty acid in the vegetable
amaranth is linoleic acid which is 49% in seeds and 46% in
leaves. Among the major saturated fatty acids, 42% linolenic
acid remains present in leaves and about 18–25% palmitic acid
in leaves, stem, and seeds. (Fernando and Bean, 1984). The
degree of unsaturation of fatty acids is over 75%.
The leaves of A. caudatus has β-sitosterol (Dixit and
Verma, 1971), while the leaves and stem of A. spinosus has
α-spinasterols and hentricontane (Banerji and Chakravarti,
1973). The roots of A. spinosus have ester of octacosanoic
acid with α-spinasterols. Massimo et al. (2004) reported βsitosterols and three major phytosterols that is, µ sitosterols,
campesterol, and stigmasterol in Amaranthus species K343,
RRC1011, K433, and K432.
Proteins
Grain Amaranth has more protein than corn and other major
cereal grains (Bejosano and Corke, 1998) (Table 5). Lysine is
the principal component which limits amino acid in cereals like
maize, wheat, and rice. EAAI value (upto 90.4%) showed that
the protein is comparable with egg protein and can be used as
a substitute for a meal (Pisarikova et al., 2005). The protein is
relatively high in sulphur containing amino acid (4.4%) (Senft,
1980), which is normally present in the pulses crops. The protein
component of amaranth is quite close to the level recommended
by the FAO/WHO for a balanced diet in humans. The protein
in grain amaranth ranges from 14.5% to 15.1% (Rodas and
Bressani, 2009) and in leaf upto 14.3 g/kg with an average of
12.4 g/kg (Shukla et al., 2003; 2006; Prakash and Pal, 1991).
The protein content in amaranth leaf is also higher than spinach,
another leafy vegetable (Table 6). The lysine in protein ranges
from 40–50 g/kg. The amino acid composition revealed that
Amaranthus is a rich source of important amino acids, namely,
alnine, valine, leucine, arginine, phenylalanine, pralines, methionines, α aminobutyric acid, tryptophan, isoleucine, and serine which suggests that amaranth is a pseudo cereal which can
be used as a substitute to nutrient cereal (Pedersen et al., 1987;
Gorinstein et al., 2002; Jaiswal et al., 1984; Misra et al., 1983;
Comparison of grain amaranthus with other grains (per 100 gms) (USDA and National Research Council)
Grain Type
Protein%
Lysine%
Carbohydrate
Calcium
Iron
Phosphorus
Amaranth
Corn
Rye
Buckwheat
Rice
Milk(human)
14.5%
0.85%
63 g
162 mg
10.0 mg
455 mg
9%
0.25%
74 g
20 mg
1.8 mg
256 mg
13%
0.40%
73 g
38 mg
2.6 mg
376 mg
12%
0.58%
72 g
33 mg
2.8 mg
282 mg
7%
0.35%
71 g
41 mg
3.3 mg
372 mg
3.5%
0.49%
5g
118 mg
Traces
93 mg
114
A. RASTOGI AND S. SHUKLA
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Table 6 Nutrients comparison of vegetable amaranth with spinach and other
leafy vegetables (per 100 g)
Components
Amaranth
Spinach
Basella
Chard
Protein
Ascorbic acid
Fiber
Carotenoids
Fat
Carbohydrate
Calcium
Iron
Potassium
3.5 g
80 mg
1.3 g
6,100 IU
0.5 g
6.5 g
267 mg
3.9 mg
411 mg
3.2 g
51 mg
0.6 g
8,100 IU
0.3 g
4.3 g
93 mg
3.1 mg
470 mg
1.8 g
102 mg
0.7 g
8,000 IU
0.3 g
3.4 g
109 mg
1.2 mg
—
2.4 g
32 mg
0.8 g
6,500 IU
0.3 g
4.6 g
88 mg
3.2 mg
550 mg
1985; Mukut and Andhiwal, 1976; Vasi and Kalintha, 1980;
Ramachandran and Phanasalkar, 1956; Jaun et al., 2007; Pisarikova et al., 2005) (Table 7). Juan et al. (2007) illustrated that
the wild species have more equilibrate amino acids which can
be introgressed in a cultivated species through hybridization.
In grain types, minerals are twice as high as the cereals
and about 66% of these minerals remain present in bran and
germ fractions (Saunders and Becker, 1984). Bran and germ
layers have high ash than the perisperm. The grains also have
high content of calcium, magnesium, iron, potassium, and zinc
(Table 7).
Vitamins
Amaranthus has necessary daily required vitamins upto a nutrient significant level and can be an excellent source for reducing vitamin deficiency (Graebner et al., 2003). In the leaves of A.
tricolor, vitamin A (carotenoids) ranged from 0.83 ± 0.02 mg/kg
and Ascorbic acid (vitamin C) ranged from 112.33 ± 5 mg/kg
(Shukla et al., 2006). Amaranth has more riboflavin (vitamin B2 )
and vitamin C than cereal and is also a good source of vitamin
E that has anti-oxidative property. An organic compound called
quercetin and Vitamin K is also reported in A. blitum (Ganju
and Puri, 1959) (Table 7).
Minerals
A large number of necessary minerals have been reported
in Amaranthus. The vegetable amaranth leaves (A. tricolor)
have potassium from 6.4 to 6.7 g/kg with an average of 3.7
± 0.26 g/kg, calcium 0.73 to 1.9 g/kg with a mean 1.7 ±
0.04 g/kg, magnesium from 2.8 to 3.0 g/kg with an average
of 2.9 ± 0.01 g/kg, zinc 434.7 to 1230 mg/kg with an average of
791.7 ± 28.98 mg/kg, iron 783–2306 mg/kg with mean 1233.8
± 50.02 mg/kg, manganese 66.7–155 mg/kg with an average
108.1 ± 3.82 mg/kg, and nickel 321.3 to 89.3 mg/kg with an
average of 222.6 ± 9.51 mg/kg (Shukla et al., 2006).
Table 7 Comparison for amino acids content (g/16 g N) of Grain amaranth
with other major cereals. (Pedersen et al., 1987)
Components
Maize
Wheat
Sorghum
Amaranth
Alanine
Arginine
Aspartic acid
Cystine
Glutamic acid
Glycine
Histidine
Isoleucine
Leucine
Lysine
Methionine
Phenyalanine
Proline
serine
Theonine
Tryptophan
Tyrosine
Valine
7.85
4.31
7.00
2.29
18.87
3.64
2.95
3.46
12.42
2.96a
1.99
4.40
9.03
4.69
3.28 c
0.70 b
3.35
4.91
3.53
4.47
4.93
2.43
30.39
3.90
2.27
3.54 c
6.73
2.75◦
1.60
4.37
9.56
4.40
2.56 b
1.33
2.51
4.46
9.22
3.64
6.48
1.91
21.01
2.93
1.97
4.05
13.67
2.03a
1.84
4.85
7.69
4.20
2.71 b
1.06
3.80
5.50
3.52
8.65
7.32
2.12
15.78
6.67
2.38
3.41
5.15 ∼
5.19
2.17
3.66
3.92
5.90
3.31 b
1.31
3.33
4.04 b
a = First limiting amino acid.
b = Second limiting amino acid.
c = Third limiting amino acid.
Fiber
Fiber is also a naturally occurring constituent of Amaranthus. In the leaves of vegetable amaranth fiber ranged between
6.95–9.65% with an average 8.39 ± 0.1% (Shukla et al., 2006).
In grain amaranth fiber is slightly lower than wheat which occurs in bran instead of perisperm layer. The fiber content ranged
from 19.5–27.9%, 35.1–49.3% and 33–44% in A cruentus, A.
hypocondriacus, and A. caudatus respectively (Pedersen et al.,
1990) .
Instead of these primary nutritionally important compounds
some other secondary metabolites that derived from metabolic
and synthetic pathways in amaranth also play an important role
in the human diet besides performing some special functions
in the plants. The descriptions of such important secondary
metabolites are discussed as follows.
Phytic Acids
The phosphorus present in the amaranth plant is produced
through the presence of phytic acid. The phytic acid ranges between 0.3 to 0.6% in Amaranthus which is equally distributed
in the seeds. It is decreased by abrasive elehulling or extraction with water. The phytic acid has a property to lower the
cholesterol level in human system.
Saponins
Saponins are found in small quantities in some of the
Amaranthus species. In seeds of A. cruentus, it varied from
0.09–0.1% of dry weight. Such a low quantity of saponins
115
NUTRITIVE VALUE OF AMARANTH
Table 8
Comparison of minerals and vitamins of grain amaranth with wheat (per 100 gm).
Grain Amaranth∗
Wheat (Anglani, 1998)
Vitamins
Grain amaranth∗
Wheat (Anglani, 1998)
Phosphorus
Potassium
Calcium
Magnesium
Iron
Zinc
Copper
455 mg
290 mg
175 mg
266 mg
17.4 mg
3.7 mg
0.77 mg
158 mg
101 mg
528 mg
266 mg
238 mg
120 mg
Ascorbic acid
Riboflavin
Folacin
Niacin
4.20 mg
0.23 mg
49 mcg
1.45 mg
Infinite
181 mg
129 mcg
∗ www.eap.mcgill.ca/CPAT
1.htm. Nov. 2008.
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Minerals
is not hazardous for consumers. The saponins present in
roots of A. spinosus are β-D-glucopyranosyl (1→4)-βD-glucopyranosyl(1→4)-β-D-glucuronopyranosyl(1→3)oleanolic acid (Banerji and Chakravarti, 1973; Banerji, 1980),
β-D-glucopyranosyl(1→2)-β-D-glucopyranosyl(1→2)-β-Dglucopyranosyl(1→3)-α-spinasterol and β-D-glucopyranosyl
(1→4)-β-D-glucopyranosyl (1→3)-α-spinasterol (Banerji,
1980). Junkuszew et al. (1998) reported four new saponins,
namely, 3-beta-(O-glucopyranosyl) ester, 3-beta-O-(alpha-Lrhamnopyranosyl (1–>3)-beta-glucuronopyranosyl)-2 beta,
3 beta, 23-trihydroxyolean-12-en-28-oic acid 28-O-(beta-Dglucopyranosyl) ester, 3 beta-O-(beta- glucopyranosyl)-2 beta,
3 beta-dihydroxy-30 norolean-12,20(29)-diene-23,28-dioic
acid 28-O-(beta-D- glucopyranosyl) ester together with known
chondrillasterol (5 alpha- stigmasta-7,22-dien-3 beta-ol) and its
3-O- glucopyranoside.
Pigments and Chlorophyll
The beautiful color of leaves in ornamental amaranth is due
to the presence of betacyanin pigments which belongs to betaline group. In amaranth betacyanin compounds are identified
as amaranthin and iso amaranthin (Dixit et al., 1991; Stintzing
et al., 2004; Repo-Carrasco-Valencia et al., 2010). The compound is O-(β –D glucopyranosuluronic acid)-5-O-β-D glucopyranosides of betadine and iso betadine respectively. The
amaranthin is an intermediate compound involved in the conversion of nitrogen compounds in the cell (Gins et al., 2002). In
the leaves of vegetable amaranth, chlorophyll a ranges between
490 ± 71.13 to 655.74 ± 109.18 and chlorophyll b ranges between 406.27 ± 65.81 to 554.92 ± 103.02 mg/kg (Shukla et al.,
2003; Ivete et al., 2004). The immense biosynthesis of amaranthine, tyrosine, and phenylalanine caused a decrease in the
contents of lignin, protein, and cellulose in leaves (Gins et al.,
2002).
Oxalates and Nitrates
Besides immense nutritional properties amaranth has two
anti-nutritional compounds, oxalates and nitrates, which ranged
from 5.1–19.2 g/kg in vegetable type and 3–16.5 g/kg in grain
types and from 0.8–0.80% mg/kg in vegetable and grain types
(Prakash and Pal, 1991; Gelinas and Seguin, 2007). The two
anti-nutritional compounds have the property to inhibit the absorption of calcium and zinc which latterly causes the development of kidney stones (Siener et al., 2006; Radek and Savage,
2008). The antinutritional compounds can easily be removed by
boiling of seeds or leaves for 5 minutes before using it for edible
purposes.
USES
As Food
Amaranthus is an important nutritional crop. It is rare plant
whose leaves are eaten as vegetable while seeds are eaten as
cereal (Oke, 1983; Saunders and Becker, 1984; Kauffman and
Hass, 1983). The spp. A. caudatus, A. hypocondriacus, and A.
cruentus used for grain purposes has tremendous potential to
increase the food production of a country. In the cultivated region of amaranth, people use amaranth as vegetable and also
as cereal in the bakery, cookies, biscuits, candies, pancakes,
pasta, and noodles formation, etc. In Peru and Bolvia, Amaranthus seeds are used as grain (Sumar, 1983) while in Mexico
the seeds of A. hypocondriacus used as grain. From popped
seeds they make candies and molasses (Early, 1977). In Peru
the seeds are popped and use as flour or bound with syrup to
make bells (Sumar, 1983). Seeds are mostly used to make laddoos in India (Vietmeyer, 1978) and sometimes it is taken with
rice after boiling with rice (Oke, 1983). In the Himalayan region, the seed flour is used to make chapattis while in Nepal
the seed flour is used as satto. In the United States, the seeds
are used to make crackers, cookies, and as cereal and also in
the preparation of baked products. Most commercial product is
the “breakfast amaranth.” A traditional use of amaranth in
Mexico and other countries is to mix popped amaranth with
honey to make a type of snack bar or snack cake. Sometimes
the whole seeds are used in a type of porridge or as a condiment
or other foods.
The amaranth is being grown for vegetable purposes throughout the tropics and Eastern Asia (Feine et al., 1979). Amaranth
leaves taste the same as spinach. For preparation of a recipe it
is recommended to boil leaves for five minutes to remove the
effect of antinutritional compounds. The cooked mass is eaten
separately or with other food (Martin and Telek, 1979). Some
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116
A. RASTOGI AND S. SHUKLA
tribal groups like Yuma, Mojava, and Cocopa cook the green
amaranth, rolled them into a ball, and dried and stored them
for winter. The Tarahumara tribe utilizes the leafy vegetable of
species A. retroflexus especially along with other leafy small
annuals. They collect the leaves during the growth cycle when
the nutrient remains at the peak in plants. In Nigeria, the leaves
are mixed with condiments for the preparation of soup (Oke,
1983). In Indonesia and Malaysia, it is commonly called bayam
and is used as a vegetable. In China the leaves and stems are
used as a frying vegetable. In the African region, the leaves
are called callaloo and used in the soup known as pepperpot
soup. In Jamaica, it is eaten routinely at breakfast and dinner.
In India, the green leaves are used directly or with other vegetables, especially potato. However, in some parts of India, grains
are used with pulses. In the Chattishgarh region, lal Bhaji is a
popular vegetable and its young stems and leaves are used as
vegetable.
As Dye
For the last 20 years, there has been an increasing trend towards the replacement of synthetic dyes by natural pigments.
For this purpose the whole plant of amaranth can be used in
preparing yellow and green dyes (Grae, 1974). The red pigment
present in the plant (part not specify) is used as a colorant in
foods and medicines (Cai et al., 2005; Cai and Corke, 1999). In
America, the roots are used to produce a dye which fades very
slightly and achieves a light yellow color with alum mordant,
tan with chrome, light olive with copper, light gold with tin,
gray with iron, and ivory with no mordant. In Bolivia and northwestern Argentina, people used red dye of Amaranthus leaves
to color alcoholic beverages while in Mexico and southwestern United States it is used for coloring maize dough (Sauer,
1950a) and for coloring foods and beverages in Ecuador (Jain
and Hauptli, 1980).
In Craft
Due to the bright red color of flowers with everlasting property, they are used in craft in some parts of world.
MEDICINAL IMPORTANCE
Classical Uses
Amaranthus being an under-utilized crop also has tremendous medicinal properties. Various species of Amaranthus have
been used in the past for the manufacturing of medicines worldwide; however, efforts still continue. In the Ayurvedic medicine
system of India, the classical text of different species of amaranth are known by different names. In Sanskrit A. spinosus
is known as Tanduliya, Meghnath, Cander, Tandulitabeej,
Vishagna, etc. Ayurveda describes the plant as diuretic, useful in cough and cold, in urinary and throat troubles, for gastric
problems and vata (Bhandari, C., 1938). The easily digestible
leaves are used in cough and cold, in swelling, and as an antidote to treat poison (Bhandari, C., 1938). A. blitum is known
as Marisa, Vaspaka, etc., in Sanskrit and is recommended as a
remedy for vata, cough and cold, and in excessive bile secretion.
The species A. caudatus is known as Rajadri, Rajagiri, etc., and
used as Pittanashka, Rucikara and also in the treatment of goiter, urinary troubles, and raktasodhaka. A. tricolor is known as
Ramasitalika and used in Raktatisara and raktapradara (Chopra
et al., 1955). Mishra (1878) describes the use of the whole plant
as a cooling agent for urinary troubles and as a lotion for external
use, in relieving pain during pregnancy, and for skin diseases.
Its roots along with amla and bark of Ashok and Daru Haldi are
used in leucorrhea.
In the Unani system of medicine, amaranth is a useful medicinal plant. The system considers the juice of the root as a remedy
for gastric problems, treating the poison of scorpion, in leucorrhea and menorrhea. The leaves are used in urinary troubles,
especially in removing kidney stones. The plant is also used to
remove skin problems (Bhandari, C., 1938). A drug “Tukhme
Ispisth” used in the system for centuries has been identified as
the seeds of A. retroflexus. According to the book “Talif Sharif,”
the plants of amaranth were used as a remedy in cough and
cold, gastric problems, blood purification, snake bite, and to
stop blood secretion from the uterus (Bhandari, C., 1938). Another important species of amaranth is A. caudatus which is also
discussed in the Indian Ayurvedic system. The whole plant is
considered as a diet for health maintenance, in blood purification, in goiter, and as a lotion (Mishra, 1978; Yoganarasimthan,
2000).
Ethanobotanical/Folk Uses
In Myth and Poetry
Amaranth or Amarant, is a name chiefly used in poetry as
a simile to express the feeling of unfading immorality. Aesop’s
fables (sixth century BC) compares the rose to Amaranth to show
the difference in fleeting and everlasting beauty. John Milton in
“Paradise Lost (1667)” and Samuel Taylor Coleridge in “Work
without Hope (1825)” also referred to Amaranth.
In India the leaves of A. caudatus are used as tea to get relief
from pulmonary problems and piles and also used in the purification of blood, stragury in scrofula, and as a diuretic (Gupta
et al. 2004). The tribals of Ghana used amaranth both medicinally and as a lotion. The Senegal tribes used boiled amaranth
roots with honey as a laxative (Anon., 1992; Kumar et al., 2004;
Nadkarni, 1954). Seeds of amaranth are used in hypertension,
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NUTRITIVE VALUE OF AMARANTH
cardiovascular diseases, reducing blood pressure, and for lowering cholesterol. In the Chattisgarh region, the traditional people
used lal bhaji for treating both internal and external diseases.
They also used amaranth for gynecological troubles, anemic
patients, dysmenorrhea, and epistaxix (Naksir). Bilaspur regional/traditional healers use lal bhaji in reducing extra fat from
the body and also as a promising herb for the patient engaged
in the obesity management program. The traditional healers of
Kondagaon region suggest that young ones wash their face with
a decoction of lalbhaji for removing pimples. The traditional
healers of Sarguja region use the fresh leaf juice in the treatment of earache.
In east Africa, the leaf known as Mchicha is prescribed by
doctors to people experiencing a low red blood cell count. In
South Africa the leaf of A. caudatus is used as an abortifacient and in pulmonary condition (Anon., 1992). The foliage of
Native American Amaranth (A. caudatus) is used to reduce hemorrhea, diarrhea, and to treat ulcerated wounds. The dried flowers are used as tea and in contraception and excessive menses.
The boiled leaves are used in swellings and in stomach upset
(Michael, 2002). In China the native amaranth roots are used
to alleviate cold, as a diuretic, and to check bleeding. The Chinese people eat amaranth during summer, believing that it can
reduce internal heat and dampness. They also use amaranth to
treat diarrhea, ulcer, inflammation, throat and mouth problems,
diabetes, and allergic reactions (Bi Fong et al., 2005). They
recommend tepid stem and leaves for washing of any affected
areas, simmered leaves with Cuttle fish for treating jaundice,
and with honey for treating dysentery. They also suggested that
amaranth leaves be used for bee stings.
In Nepal, the juice of amaranth root is used to treat fever,
urinary trouble, diarrhea menorrhagia, gonorrhea, eczema, colic
lactogogue, and dysentery and its combination with the juice of
Dichrophela integra and Rubus elliptices in stomach disorders.
Boiled roots and leaves are given to children as laxative and
emollient and as poultice in abscesses, in burns, and snake bite
(Ministry of Forest (Nepal), 1970).
In Swaziland and South Africa, the ash of the plant is used either alone or mixed with an equal quantity of powdered tobacco
for making snuff (Edward, 1978).
Another species, A. blitum of Amaranthus, is also used medically. It is a cooling, stomachic, emollient and slightly astringent,
useful in biliousness, hemorrhea, and diathesis. It is also used in
the preparation of tonic and remediation of round worm (Anon.,
1992; Jain and Defillips, 1991). In the Garhwal region, the leaf
of A. hybridus is used for diarrhea and leucorrhea and the root in
gonorrohea, colic, and eczema (Rajwar, 1983). The whole plant
of A. hybridus subsp. cruentus (Linn.) is an astringent (Jain and
Defillips, 1991) used in piles to purify blood, as a diuretic, and
an antiscorbutic (Anon., 1992). The plant of subsp. A. hybridus
is used as a detergent and astringent, for menorrhagia, diarrhea,
dysentery, ulcerated condition of throat and mouth, leucorrhea,
and for washing of ulcers and sores, etc. (Anon., 1992). Another
important species, A. spinosus, has been used for a long time as
an important medicinal plant. The whole plant is recommended
117
as an enmenagogue and galactagogue. The plant is used as a
refrigent, diuretic, purgative (Sebasttian and Bhandari, 1984),
as a laxative in vomiting (Jain and Defillips, 1991), enemas
in stomach troubles, against inflammation, in uterine bleeding,
as an antidote to snake bite, in wounds, as wormicidal, for
reduction of excessive bleeding during menstruation, in skin
diseases, and cough and cold (Anon., 1992; Suresh et al., 1995;
Bhatnagar et al., 1973; Srivastava et al., 1866; Raj and Patel,
1978; Joshi et al., 1980; Banerjee and Banerjee, 1986; Basak,
1997; Kumar and Goel, 1998; Sudhakar and Chetty, 1998; Jain
and Defillips, 1991; Yoganarasimthan, 2000; Ministry of Forest
(Nepal), 1970; Vaidyaratnam, 1993; Lindley, 1981; Prajapati
et al., 2003). The roots are thermogenic and hemostatic and
useful in vitiated condition of Kalpa, menorrhea, hemoptysis,
hematemensis, leucorrhea, and anti-diarrheal (Sebasttian and
Bhandari, 1984; Mohanty et al., 1996), an emollient (Singh
et al., 1998), an anti-spasmodic (Bhatnagar et al., 1973), and in
ear diseases (Reddy et al., 1989). The plant is also considered
as sudorific and febrifuge and recommended for eruptive fevers
(Yoganarasimthan, 2000). The decoction of the plant improves
digestion, treats kidney problems, and is used as mouthwash for
toothache. The decoction with palm nut soap is used to arrest
miscarriage (Bakshi et al., 1992).
Siddha used the plant in its entirety for treating snake
bite, amenorrhoea, leucorrhoea, and abscesses. Roots are used
in uterine diseases and burning sensation (Yoganarasimthan,
2000). The Lodha tribes use smoke of the root as a hallucinogen and eat root paste; however, this can cause insanity (Bakshi
et al., 1992). The tribes of Medinipur tie a piece (2 cm) of root in
black thread on left arm of a pregnant lady to cure piles (Bakshi
et al., 1992). In Ivory Coast the plant is used for treating leprosy
(Anon., 1992).
The species A. tricolor is also used medically. The plant is
used as an astringent and diuretic, in menorrhea, diarrhea, dysentery, cuts and wounds, cough and bronchitis, paralysis in cattle,
and as a poultice to wash wounds (Tripathi et al., 1996; Kapoor
and Kapoor, 1980; Srivastava et al., 1980; Singh and Dhar, 1993;
Singh et al., 1998; Jain and Defillips, 1991). The leaves are used
in jaundice, intestinal and urinary discharges, menorrhea, fever,
blood disorder, piles, and strangury scrogfula (Borthakur and
Goswami, 1995; Gupta et al., 2004; Sankarnarayanan, 1988;
Tripathi et al., 1996; Yoganarasimthan, 2000). The roots are
used in gonorrhea, eczema, and as a demulcent. Its decoction
is used for stricture, piles, and diarrhea. The roots and seeds
are also used to cure leucorrhea and impotency (Chopra et al.,
1956; Anon., 1992; Prajapati et al., 2003; Bakshi et al., 1992). In
present times the poultice is used in ulcerated throat, toothache,
and as an astringent and cooling agent (Bakshi et al., 1992).
Other species, A. viridis, has anthelmintic properties and is
also used as a laxative. It is used in inflammation (Tripathi
et al., 1996; Kakrani and Saluja, 1994) in snake bite, and scorpion stings (Jain and Defillips, 1991). The roots of the plant
are recommended in eczema and against ring worm (Siddiqui
et al., 1989). Traditionally, Lodha tribes used root paste on burns
(Bakshi et al., 1992). In Brazil the plant is used as a diuretic and
118
A. RASTOGI AND S. SHUKLA
galactagogue. The leaves are used as a poultice in inflammation,
boils, and abscesses (Anon., 1992). The species A. oleracea is
used for emollients, cooling stomach, hemorrhea, diathesis, and
biliousness (Jain and Defillips, 1991). The species A. polygonam is used as a demulcent, aphrodisiac, and in impotency. The
decoction of the roots and leaves are used to treat leucorrhea,
menorrhea, and diarrhea. The leaves are used as a poultice in
inflammation, painful body parts, and abscesses. The roots are
recommended for colic gonorrhea and eczema.
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BIOLOGICAL EFFECTS
Though Amaranthus species can be used for different beneficial purposes but some weedy species have negative as well
as allergic effects. Suma et al. (2002) studied the allelophatic
effect of weedy amaranths viz. A. retroflexus, A. spinosus, and A.
viridis. They reported that these species inhibit crop growth and
characterizes allelochemicals from A. spinosus. The pollen of A.
spinosus causes pollen allergic diseases and pollen of A. viridis
are found to be associated with skin diseases. Petroleum ether
extract of aerial parts of A. tricolor showed antibacterial activity against gram positive bacteria Staphylococcus aurens, Staph
albus, and Streptococcus viridians (Sharma and Mukat, 1991).
The seeds of A. blitum growing in dung revealed in vitro antibacterial activity against Pseudomonas cichorii and Xanthomonas
campestris and A. viridis seed inhibits bacteria Bacillus subtilis
and E. coli MLS-16 (Bagchi et al., 1997; 1999). The extract of
leaves of A. viridis showed strong nematocidal activity against
second larvae of Meloidogyne incognita in vitro at various concentrations (Nandal and Bhatti, 1983; Siddiqui and Alam, 1997).
Amoli et al. (2009) determined the total toxicity of Amaranthus
retroflexus L. The plant extract has been tested for bioactivity in
Artemia salina and cytotoxicity against bovine kidney for cells.
The bovine kidney cells were exposed to various concentrations
of the plant extracts (100 ppm-0.1 ppm). After treating with 100
and 0.1 ppm for 24 h, the viability of the cells were reduced
by about 49% and 35%, respectively, in MTT viability assay.
The study confirmed that Amaranthus retroflexus has a cytotoxic
effect and more specifically to renal cells.
MOLECULAR APPROACHES
A lot of molecular work has been carried out by various
workers on Amaranth to study the basic structure of different
molecules and genes responsible for various important proteins,
starch and other compounds present in it. Raina and Dutta (1992)
isolated a well-balanced albumin protein (35Kda) with four isoforms from the seeds of A. hypocondriacus. Chakraborty et al.
(2000) expressed a non-allergic seed albumin AmA1 with a well
balanced amino acid composition encoded from a single gene
of A. hypocondricus using granule bound starch synthatase enzyme (GBSS) and cauliflower mosaic virus 35S promoter to
express in potato to increase the nutritive value of potato. Mar-
cone et al. (1994) purified other major albumin fraction (i.e.,
albumin 1) of A. hypocondriacus having a molecular mass of
133400d. The secondary, tertiary, and quaternary structure of
storage globulin protein were also studied through electron microscope which showed that it had some overall structure as
a dicot plant (Marcone et al., 1994). Similarly, RomeroZepeda
et al. (1996) isolated IIs amaranth seed globulin with molecular
weight 389 kda from A. hypocondricus and termed it Amaranthin. Dela Rosa (1996) extracted a globulin protein of molecular
weight 398 kda from amaranth seeds and purified it by gel filtration and ultracentrifuge. The deduced amino acid sequence
confirmed that protein synthesized was similar to other IIs like
protein and also had the same ancestral gene compared to the dicot and the monocot species. Castellani et al. (1999) studied the
role of disulphide bands on the structure stability of amaranth
globule protein and noticed that its polymerization was high
(51+/-1mu mol/g) and similar to soybean IIs globulin content.
Ramirez-Medeles et al. (2003) also isolated and purified nsLTP
protein with molecular mass of 9747.29 Da from the seeds of
A. hypocondriacus by gel filtration and reverse phase HPLC.
The protein had a alpha helical structure typical to other plants
nsLTP1 and had 40 to 57% sequences identical to other plants.
Lozanov et al. (1997) reported total synthesis of a-amylase inhibitor, a 32 residue long peptide with 3 disulphide bridges
from the seeds of Amaranthus. Aphalo et al. (2004) studied the
surface characteristic and assess the homology of globulin protein, the polymerized 11s amaranth globulin with other storage
proteins which showed that globulin P unitary molecules with
aggregates had a similar reactive surface. Chen et al. (2004) separated ribosome inactivating protein (molecular weight 29kda)
from the seeds of Amaranthus mangostanus, a kind of vegetable
which had an isoelectric point greater than 9. This protein can be
used for medicinal purposes which can be applied in the preparation of antitumor, antiviral, and anti HIV drug formation. For
the first time Petruccelli et al. (2007) studied the role of short
sequence conserved among 7S and 11S proteins in vacuolor
sorting and interaction among proteins of different classes in A.
hypocondriacus. Legaria et al. (1998) isolated a genome clone
(a hybadh 4) and a clone (a hybadh A), both encoded for betaine aldehyde dehydrogenase from A. hypocondriacus. Molina
et al. (2008) reported that the seeds of Amaranthus also contain
non-processed 11 s precursors which are constituent of albumin
2.
Two antimicrobial polypeptides were isolated from A. caudatus seeds (Ac AMP 1 and Ac-AMP 2) and had a molecular
mass of 3025 and 3181da with PI values 9 and 10 (Broekaert
et al., 1992). The amino acid sequence was identical in both
polypeptides except the latter had one additional residue at the
carboxyl terminus. The sequences had high homology to other
chitin binding protein. They could inhibit the growth of different plant pathogen fungi at much low doses and other known
fungal chitin binding proteins. They also show activity to gram
positive bacteria. Rodriguez (1993) purified a 69 amino acids
protein with high catene and valine, arginine, glutamic acid
and lack in methionine protein protinase inhibitor of molecular
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NUTRITIVE VALUE OF AMARANTH
weight 7400 having an isoelectric point 7.5 from a seed extract of A. hypocondriacus. It had the tendency to inhibit the
activity of larvae of insect Prostephanus truncates. Another major trypsin inhibitor (AMTI) which is of 69 amino acid protein
had the homology with members of the potato I inhibitor family, which were isolated and sequenced by Valdes-Rodriguez in
1999. He cloned and expressed 394 bp cDNA sequences with
an open reading frame corresponding to a polypeptide of 76
amino acids residues and encoded these in various vegetative
tissues of amaranth plant during seed development and imbibitions. Further, Rodriguez et al. (2007) encoded and characterized
Cysteine proteases inhibitor (Al CPI) which had a polypeptide
of 247 amino acids including a putative n-terminal signal peptide. The sequence also had G and PW conserved as motifs, to
conserve LARFAV sequence for phytocystatins, and the relative site QWAG. The sequence (ALCPI) showed a significant
homology to other plant cystatins. Its expression analyses indicated that it expressed in mature seeds and decreased during
germination which described that single cystatin protein could
have a regulatory role in germination. Similarly, Tamir et al.
(1996) isolated a 8 kd trypsin chromatrypsin inhibitor proteins
from seeds of A. hypocondriacus through affinity chromatography on trypsin sepharose and by HPLC. The protein was stable at neutral and alkaline pH and was relatively thermostable.
Pribylova et al. (2006) tried to develop an easy and cheap PCR
method for detection of an antimicrobial peptide from Amaranthus for transformation of proteins. Again, Pribylova et al.
(2008) used the primer to amplify the gene AMP2 in A. caudatus. He noticed that the same genes were present in other seven
species of Amaranth, namely A. albus, A. cruentus, A. hybridus,
A. hypocondriacus, A. retroflexus, and A. tricolor and observed
that all sequences were identical except for two polymorphisms
and this polymorphism was present outside the region encoding
the chitin binding peptide domain, thus not affecting the proper
functioning of the antimicrobial peptide.
Boinski et al. (1993) studied the post-transcription control of
cell type specific gene expression in bundle sheath and mesophyll chloroplast of A. hypocondriacus and demonstrated that
the rubisco LSU polypeptide was present only in chloroplast
preparation of bundle sheath cells and Pyruvate orthophosphate
dikinase (PPDK). A nuclear encoded chloroplastic enzyme was
only found in a mesophyll chloroplast preparation. Brery et al.
(1997) studied photosynthetic gene expression in developing
the leaves of a NAD-ME type C4 dicot in amaranth and showed
that the C4 gene independently regulated by a different multiple control mechanism instead of environmental and metabolic
signals. Ptushenko et al. (2002) tried to study the interaction of
Amaranthin protein to ETC of chloroplasts at the PS level. They
found that the protein affect ETC near PS I and had no significant inhibitory effect on the light dependent formation of the
transmembrane PH gradient at the same time. Long et al. (1994)
purified mitochondrial NAD dependent malic enzyme made up
of 2 subunits, 65 and 60 kd designated α and β, respectively,
to catalyze the decarboxylation of 4 carbon malate in bundle
sheath cells releasing CO2 for the calvin cycle. The Zummogold
119
electromicroscopy using α subunit antiserum demonstrated that
the protein was localized specifically in mitochondrial matrix
bundle sheath cells and had similarity with the other plants,
animals, and bacterial malic enzyme. Barrio and Anon (2010)
isolated protein from the seeds of Amaranthus mantegazzianus
that exhibit potential of antitumor properties using four different
tumor-derived and in vitro-transformed cell lines with different
morphology and tumorigenicity (MC3T3E1, UMR106, Caco-2,
and TC7). The MPI showed an antiproliferative effect on four
cell lines with different potencies. The MPI produced morphological changes and caused a rearrangement of the cytoskeleton
in UMR106 cell line.
Praznik et al. (1999) for the first time studied the molecular background of technological properties, for example, gelatinization, stability to mechanical stress, resistance to stability
in continuous freeze cycle of selected starch of maize, amaranth,
C. quinoa, etc. and correlated these properties with molecular
features such as branching characteristic in terms of iodine complexing potential molar mass occupied glucan volume, packing
density of glucan oils, and theological properties.
Bello Pezec (1998) studied the macromolecular feature of
starch with the help of high performance size exclusion chromatography (HPSEC), static light scattering (SCS), and dynamic light scattering (DLS) techniques for characterization of
amaranth starch and showed the sphere and the globular structure of starch. Bunzel et al. (2005) studied the association of
non-starch polysaccharides and ferulic acid in dietary fiber of
grain amaranth (A. caudatus). They reported that through the
ferulocylated oligosaccharides ferulic acid was predominately
bounded to pectin arabians and galactins in amaranth insoluble fiber. The compound 50 transferoloyl L arabinofuranose
was isolated in pure form from pectic arabinans but also from
arabinoxylans. Park et al. (2009) isolated and characterized a
full-length cDNA clone encoding granule-bound starch synthase I (GBSSI = Waxy gene) from grain amaranth (Amaranthus cruentus L.) perisperm. Segregation of amylose content in
F2 population suggested that the amylose content of A. cruentus is controlled by a single gene. cDNA clone of this gene is
2076 bp in length and contains an open reading frame of 1821 bp
corresponding to a polypeptide of 606 amino acids residues, including a transit peptide of 77 amino acids. Sarkar et al. (2009)
isolated a water-soluble polysaccharide (PS-I), from the aqueous extract of the stems of Amaranthus tricolor Linn. (Amaranthus gangeticus L.), was found to consist of L-arabinose,
methyl-D-galacturonate, D-galactose, and 3-O-Ac-L-rhamnose
in a molar ratio of nearly 1:1:1:1. Kong et al. (2009) determined
the fine structure of amylopectin from grain amaranth. Amaranthus amylopectin was hydrolyzed with alpha-amylase, and
single clusters and a group of clusters (domain) were isolated
by methanol precipitation and showed that the domain fraction
contained approximately 2.2 clusters, and single clusters composed of approximately 13 chains. The phi,beta-limit dextrins
of the clusters were further hydrolyzed with alpha-amylase to
characterize their building block composition. The average DP
of the branched blocks was approximately 11 and contained on
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A. RASTOGI AND S. SHUKLA
an average of approximately 2.5 chains. Their average chain
length, internal chain length, and degree of branching were approximately 4.3, 2.8, and 14, respectively.
To study the genetic diversity based on molecular markers
among the Amaranthus several efforts had been made. Sun et al.
(1999) used a low cost DNA probe obtained by isolation of various classes of repetitive DNA sequences, including satellite,
mini satellite, rDNA, retro transposones like sequences, and
other unidentical novel repetitive sequences to study genetic diversity among 24 cultivated and wild species of Amaranthus.
DNA fingerprinting by repetitive DNA probes revealed a different level of polymorphism in Amaranth genome. Wetzel et al.
(1999) used the PCR based molecule identification system to
distinguish the weedy species of genus Amaranth. Lee et al.
(2008) isolated 12 polymorphic microsatellite markers from 20
accessions of A. hypocondriacus and reported 92 alleles with
average alleles 7.7 per locus with heterozygosity value ranged
from 0.0 to 0.95. The results demonstrated that it has wide applicability of markers for the study of intra and inter specific
genetic diversity as well as evolutionary relationships among
cultivated and wild amaranth. Mallory et al. (2008) developed
microsatellite enriched libraries by sequencing 1457 clones in
grain amaranth and reported that 179 microsatellites were polymorphic across accessions of three grain amaranth. In polymorphic satellites a total of 731 alleles were reported with an
average of 4 alleles per locus which suggested that 3 cultivated
species were evolved from different A. hybridus ancestors.
BREEDING APPROACHES
Amaranthus is a rich and cheap source of daily required nutrients as already mentioned previously. In recent years it is
gaining importance as an alternative crop to other important cereals. To meet out increasing global demand and simultaneously
to decrease pressure on major cereal crops, the development of
high yielding varieties rich in nutrients are very important. In
this direction various studies and breeding efforts have been initiated worldwide, especially in India. In the development of a
high yielding variety, the presence of genetic diversity among
available germplasm lines plays a major role (Shukla et al.,
2010). The extent of genetic diversity among different lines of
three species of grain Amaranthus and among 66 strains of vegetable amaranth have been explored by Gupta and Gudu (1991),
Shukla et al. (2002), and Pandey (2009), respectively. Transue
et al. (1994); Chan and Sun (1997), Ranade et al. (1997), Xu
and Sun (2001), Stefunova and Bezo (2003), and Costea et al.
(2006) studied genetic diversity among different Amaranthus
germplasm based on RAPD or AFLP markers or isozymes.
Overall study showed that A. hypocondriacus and A. caudatus
were more genetically similar than A. cruentus, putative origin of A. dubious was from A. hybridus and A. spinosus, A.
caudatus, and A. tricolor were however, intermediate. A. hybridus and A. dubious were more similar. Ray and Roy (2007)
for the first time made phylogenetic relationships among the
members of amaranthaceae and chenopodiaceae. For improvement of cultivars not only the knowledge of genetic diversity
is important but also knowledge of magnitude of variation in
the available germplasms, interdependence of quantitative characters with yield, extent of environmental influence on these
factors, heritability, but genetic advance of various contributing
traits are also required. Keeping this in mind various workers
made their efforts in this direction as well. Imeri et al. (1987),
George et al. (1989), and Shukla and Singh (2003) studied the
genetic variability and correlation among the different varieties
of grain amaranth. Similarly, Reddy and Varalakshmi (1994),
Shukla and Singh (2000), and Shukla et al. (2004; 2005; 2006;
2006) studied genetic variability among the available accessions
and also studied different genetic parameters for different traits
over different cuttings in vegetable Amaranthus and suggested
that the enhancement of their foliage yield and its quality traits
could be achieved through selection of plant types based on
all of its component traits. Similarly, Prakash and Pal (1991);
Shukla et al. (2000; 2002; 2004), Shukla and Singh (2000),
and Bhargava et al. (2004) studied the correlation among foliage yield and its contributing traits in vegetable amaranth over
different cuttings. However, Katiyar et al. (2000), Reddy and
Varalakshmi (1994), and Shukla and Singh (2001; 2003) studied the correlation over different traits. Katiyar et al. (2000) and
Mathe-Gaspar (2001) isolated lines of grain amaranth suitable
in sodic soil and tolerant to drought. They reported that grain
amaranths due to drought tolerance ability, can be suitable and
economically feasible commercial crop for wastelands. MatheGaspar (2001) characterized the important phenological traits
of Amaranthus cultivation. Similarly, Escudero et al. (1999) in
A. muricatus, Lakshmi and Vimala (2000) in A. gangeticus and
Shukla et al. (2003; 2006) in A. tricolor made their efforts to
study the nutritional and mineral components. Previously, Rangarajan et al. (1998) also evaluated the grain and leafy amaranth
for iron and found that amaranth has relatively much iron than
other leafy vegetable. Sharma et al. (2001) studied the stability
of grain amaranth genotypes over different years and noted significant genotype x environment interactions for all the traits.
Similarly, Varalakshmi (2003) and Shukla and Singh (2003)
studied the stability for economic traits in vegetable amaranth.
Lehmann et al. (1991) and Reddy and Varalakshmi (1998) studied the combining ability and heterosis in amaranthus. Wu et al.
(2000) evaluated the Amaranthus collection of China and concluded that many species were sensitive to day length. Cultivated species had high grain yield but were more sensitive to
diseases. Jamriska (1993) and Henderson et al. (2000) studied
row spacing, plant population, and cultivar effects on grain and
found that yield depends upon weather conditions and narrow
row spacing produced average higher yields than the stands
with wider row spacing. Teutonico and Knorr (1984) studied the antinutritional components of A. tricolor through tissue culture. Later on, Bennici et al. (1992; 1997) also made
tissue culture studies in different lines of grain Amaranthus
species. They suggested that genotype, growth regulator dose,
type, and physiological stage of explants were important factors
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NUTRITIVE VALUE OF AMARANTH
for Amaranthus in in vitro regeneration. Law-Ogbomo and Ajayi
(2009) determined the influence of planting density and poultry manure application on the growth and yield of Amaranthus cruentus (Linnaeus) and reported that the combination of
62,500 plants per hectare and application of poultry manure of
12 t/ha, provided the highest yield (15.74 t/ha). Khandaker et al.
(2010) evaluated biomass yield and accumulation of betacyanin,
chlorophyll, total polyphenol, and antioxidant activity in Amaranthus tricolor L. under five shades made up of white, blue,
green, yellow, and black polythene and a control. They reported
that blue polythene has the potential to increase the yield with
health beneficiary bioactive compounds betacyanins, polyphenol, and antioxidant activity during the low temperature regime
in the spring season.
FUTURE PROSPECTS
The overview of available literature showed that Amaranthus
possesses versatility in its nature and uses. The presence of
genetic diversity among lines/species of both grain and vegetable types opened the path for breeders for its improvement
and development of nutritionally rich varieties by applying
conventional breeding approaches. The presence of a large
amount of different amino acids in seed embryo provides
the way for the development of specific nutritionally rich
varieties (Tomoskozi et al., 2009). The species which have
drought tolerance and capability to grow in sodic soil can be
explored for wastelands cultivation. Amaranth seed oil contains
non-polar lipids with a high degree of unsaturation still needed
to be identified followed by the establishment of accurate
oil refining processes. At the present time the development
of varieties with low/nil oxalate and nitrate components is
required which needs extensive research. Teutonico and Knorr
(1984) obtained reduction in oxalate and nitrate through tissue
culture techniques, thereby opening the possibilities for the
development of such varieties by tissue culture. Demko (1997)
noticed the use of Amaranthus retroflexus as a new alternative
of renewable energy resources and found that it reached 75 to
90% of the calorific power of wood. So, the efforts should be
made to explore other species also for the purpose of renewable
resources. Based on available information on medicinal uses of
Amaranthus for pharmaceutical industries, researchers should
focus on separation and characterization of compounds having
medicinal properties. Thus, Amaranthus can be a future crop for
diversified purposes and can solve the problem of malnutrition,
especially for developing countries.
REFERENCES
Allen, P. (1961). Die Amaranthaceen Mittleleuropas Carl Verlag, Munchen.
Anonymous (1992). The Wealth of India. Publications and Information Directorate, CSIR, New Delhi, India.
Anglani, C. (1998). Wheat minerals: A review. Plants Food Human Nut. 52:
177–186.
121
Aphalo, P., Castellan, O. F., Martinez, E. N., and Anon, M. C. (2004). Surface
physiological properties of globulin P amaranth protein. J Agric Food Chem.
52: 616–622.
Amoli, J. S., Sadighara, P., Barin, A., Yazdani, A., and Satari, S. (2009). Biological screening of Amaranthus retroflexus L. (Amaranthaceae). Brazilian J
Pharma. 19: 617–620.
Badami, R. C. and Patil, K. B. (1976). Studies on vegetable oils of the genus
Amaranthus. J Oil Technol Assoc, India. 8: 131–134.
Bagchi, G. D., Singh, A., Khanuja, S. P. S., Singh, S. C., Bansal, R. P., and
Kumar, S. (1997). Antibacterial activities observed in the seeds of higher
plants found to grow on the cattle dung. JMAPS. 19: 980–987.
Bagchi, G. D., Singh, A., Khanuja, S. P. S., Singh, S. C., Bansal, R. P., and
Kumar, S. (1999). Wide spectrum antibacterial and antifungal activities in
the seeds of some coprophilous plants of North Indian plains. J Ethnopharm.
64: 69–77.
Bakshi, D. N. G., Sensarma, P., and Pal, D. C. (1999). A Lexicon of Medicinal
Plants of India, Naya Prakash, Calcutta, India.
Banerji, N. (1980). Two new saponins from the root of Amaranthus spinosus
Linn. J Indian Chem Soc. 57: 417–419.
Banerji, N. and Chakravarti, R. N. (1973). Constituents of Amaranthus spinosus
Linn. (Amaranthaceae). J Inst Chem (India). 45: 205–206.
Banerjee, A. K. and Banerjee, I. (1986). A survey of the medicinal plants in
Shevaroy hills. J Econ Tox Bot. 8: 271–290.
Basak, S. K. (1997). Medicinal plants of Bankura (W.B.) and their uses. J Nat
Bot Soc. 51: 61–68.
Becker, R., Wheeler, E. L., Lorenz, K., Stafford, A. E., Grosjean, O. K.,
Betschart, A., and Saunders, R. M. (1981). A compositional study of Amaranth grain. J Food Sci. 46: 1175.
Bejosano, F. P. and Corke, H. (1998). Protein quality evaluation of Amaranthus
wholemeal flours and protein concentrates. J Sci Food Agric. 76: 100–106.
Bello Pezec, L. A., de Leon, Y. P., Agama-Acevedo, E., and Parades-Lopez,
O. (1998). Isolation and partial characterization of amaranth and banana
starches. Starch Strake. 50: 409–413.
Bennici, A., Schiff, S., and Bovelli, R. (1992). Invitro culture of species and
varieties of 4 Amaranthus species. Euphytica. 62: 181–186.
Bennici, A., Grifoni, T., Schiff, S., and Bovelli, R. (1997). Studies on callus
growth and morphogenesis in several species and lines in Amaranthus. Plant
Cell Tissue Organ Cult. 49: 29–33.
Brery, J. O., McCormac, D. J., Long, J. J., Boinski, J., and Corey, A. C. (1997).
Photosynthetic gene expression in amaranth an NAD-MEtype C-4 dicot. Aus
J Plant Physio. 24: 423–428.
Bhandari, C. (1938). Vanaushadhi Chandrodaya: An Encyclopedia of Indian
Botanics and Herbs. Chowkambha Sanskrit Series, Varanasi, U.P., India.
Bhargava, A., Shukla, S., Chatterjee, A., and Singh, S. P. (2004). Selection
response in vegetable amaranth (A. tricolor) for different foliage cuttings. J
Applied Horticulturae. 6: 43–44.
Bhatnagar, L. S., Singh, V. K., and Pandey, G. (1973). Medico-botanical studies
on the flora of Ghatigaon Forests, Gwalior, and Madhya Pradesh. J Res Ind
Med. 8: 62–100.
Bi Fong, L., Bor Luen, C., and Jin Yuarn, L. (2005). Amaranthus spinosus
water extract directly stimulates proliferation of β- lymphocytes in vitro.
International Immuno Pharma. 5: 711–722.
Boinski, J. J., Wang, J. L., Xu, P., Hotchkiss, T., and Berry, J. O. (1993).
Posttrancriptional control of cell type specific gene expression in bundle
sheath and mesophyll chloroplasts of Amaranthus hypocondriacus. Plant
Mol Biol. 22: 397–410.
Borthakur, S. K. and Goswami, N. (1995). Herbal remedies from Dimoria of
Kamrup district of assam in Northeastern India. Fitoterapia. 66: 333–340.
Bostid, D. (1984) Amaranth Modern Prospects for an Ancient Crop. 74 p.
Bunzel, M., Ralph, J., and Steinhart, H. (2005). Assocation of non starch
polysaccharides and ferculic acid in grain amaranth (Amaranthus caudatus
L.) dietary fiber. Mol Nut Food Res. 49: 551–559.
Broekaert, W. F., Marien, W., Terras, F. R. G., De Bolle, M. F. C., Damme, P.
P., Jozef, V., Dillen, L., Claeys, M., and Rees, S. B. (1992). Antimicrobial
peptides from Amaranthus caudatus seeds with sequence homology to the
cysteine/glycine-rich domain of chitin-binding proteins. Biochemistry. 31:
4308–4314.
Downloaded by [National Botanical Research Inst] at 04:19 22 February 2013
122
A. RASTOGI AND S. SHUKLA
Barrio, D. A. and Anon, M. C. (2010). Potential antitumor properties of a protein
isolate obtained from the seeds of Amaranthus mantegazzianus. Eur J Nutr.
49: 73–82.
Cai, Y. and Corke, H. (1999). Amaranthus betacyanin pigmants applied model
food systems. J Food Sci. 64: 869–873.
Cai, Y., Sun, M., and Corke, H. (2005). Characterization and application of
Betalain pigmaent from plants of the Amaranthaceae. Trends Food Sci Tech.
16: 370–376.
Castellani, O. F., Martinez, E. N., and Anon, M. C. (1999). Role of disulphide
bonds upon the structural stability of an amaranth globulin. J Agric Food
Chem. 47: 3001–3008.
Chan, K. F. and Sun, M. (1997). Genetic diversity and relationships detected by
isozyme and RAPD analysis of crop and wild species Amaranthus. TAG. 95:
865–873.
Chaturvedi, A., Sarojni, G., and Devi, N. L. (1993). Hypocholesterolemic effect
of Amaranth seeds (Amaranthus esculentus). Plants Food Human Nut. 44:
63–70.
Chen, M., Wang, Y., and Wang, Z. (2004). Amaranth seed ribosome inactivating protein. Fujian Research Institute Matter Structure. Chinese Patent No.
2004–488613.
Chidambaram, N. and Iyer, R. R. (1945). Chemical examination of the seeds of
Amaranthus gangeticus. Part 1. The fatty oil from the seeds. J Indian Chem
Soc. 22: 117–118.
Chopra, R. N., Colonel, B. T., and Chopra, I. C. (1955). A Review of Work
on Indian Medicinal Plants. Indian Council of Medical Research, Cambridge
Printing Work, New Delhi.
Chopra, R. N., Nayar, S. C., and Chopra, I. C. (1956). Glossary of Indian
Medicinal Plants. CSIR, New Delhi.
Chopra, R. N., Nayar, S. L., and Chopra, I. C. (1986). Glossary of Indian
Medicinal Plants. Council of Scientific and Industrial Research, New Delhi.
Chakraborty, S., Chakraborty, N., and Datta, A. (2000). Increased nutritive value
of transgenic potato by expressing a non allergic seed albumin gene from A.
hypocondriacus. PNAS. 97: 3724–3729.
Coimbra, S. and Salema, R. (1994). Amaranthus hypochondriacus-seed structure and localization of seed reserves. Annals Bot. 74: 373–379.
Costea, M., Brenner, D. M., Tardif, F. J., Tan, Y. F., and Sun, M. (2006). Delimitation of Amaranthus cruentus L. and Amaranthus caudatus L. using
micromorphology and AFLP analysis: An application in germplasm identification. Gen Res Crop Evol. 53: 1625–1633.
Dela Rosa, A. P. B., HerreaEstrella, A., Utsumi, S., and ParedesLopez, O.
(1996). Molecular characterization, cloning and structural analysis of a cDNA
encoding an amaranth globulin. J Plant Physio. 149: 527–532.
Demko, J. (1997). Analysis of energy of Amaranthus retroflexus. Rostlinna
Vyroba. 53: 65–68.
Dixit, B. S., Srivasatava, S. N., and Pal, M. (1991). Pigments of Amaranthus,
Celosia, Beta vulgaris and their utilization. Indian J Nat Prod. 7: 12–14.
Dixit, V. K. and Verma, K. C. (1971). Some observations on fixed oil of seeds
of Amaranthus caudatus Linn. Indian Oil Soap J. 36: 273–274.
Early, D. K. (1977). Cultivation and uses of Amaranth in contemporary Mexico.
Proceedings of the first Amaranth Seminar, p. 39. Rodale Press, Emmaus,
PA.
Edward, S. A. (1978). Medicinal Plants of South Africa. Reference Publication
Inc., Michigan.
Escudero, N. L., Albarracin, G., Fernandez, S., de Arellano, L. M., and Mucciarelli, S. (1999). Nutrient and antinutrient composition of Amaranthus muricatus. Plant Food Human Nutr. 54: 327–336.
Feine, L. B., Harwood, R. R., Kauffman, C. S., and Sneft, J. P. (1979). Amaranth:
Gentle giant of the past and future. In: New Agricultural Crops, p. 105. Rodale
Press, Emmaus, PA.
Fernando, T. and Bean, G. (1984). Fatty acids and sterols in Amaranthus tricolor
L. Food Chem. 15: 233–237.
Franseen, A. S., Skinner, D. Z., Al-Khatib, and Horak, M. J. (2001). Pollen
morphological differences in Amaranthus species and interspecific hybrids.
Weed Sci. 49(6): 732–737.
Ganju, K. and Puri, B. (1959). Bioflavanoids from Indian vegetables and fruits.
Indian J Med Res. 45: 197–200.
Gamel, T. H., Mesallam, A. S., Damir, A. A., Shekib, L. A., and Linssen, J. P.
(2007). Characterization of Amaranth seed oils. J Food Lipids. 14: 323–334.
Gelinas, B. and Seguin, P. (2007). Oxalates in grain Amaranth. J Agric Food
Chem. 55: 4789–4794.
George, S., Barat, G. K., Sivakamin, N., and Choudhury, B. (1989). Source and
variability for nutritive aspects in amaranth (Amaranthus species). Indian J
Agric Sci. 59: 274–275.
Graebner, I. T., Siqueira, E. M. A., Arruda, S. F., and Souza, E. M. T. (2004).
Carotenoids from native Brazalian dark green vagetables and bioavailable: A
study in rats. Nutr Res. 24: 671–679.
Gins, M. S., Gins, V. K., and Kononkov, P. F. (2002). Change in the biochemical
composition of Amaranth leaves during selection for increased amaranthine
content. Appl Biochem Microbio. 38: 474–479.
Gorinstein, S., Pawelzik, E., Delgado-Licon, E., Haruenkit, R., Weisz, M.,
and Trakhtenberg, S. (2002). Characterization of Pseudocereal and cereal proteins by protein and amino acid analyses. J Sci Food Agric. 82:
886–891.
Goering, K. J. (1967). New Starches. II. The properties of starch chunks from
Amaranthus retroflexus. Cereal Chem. 44: 245–252.
Grae, I. (1974). Nature’s Colors-Dyes from Plants. MacMillan, New York.
Grazdiene, D. (2007). Chemical composition and properties of oil from Amaranthus seed growing in Litherania. Verterinarija IR Zootechnika. 39: 22–29.
Grubben, G. J. H. and Van Stolen, D. H. (1981). Genetic resources and Amaranths. International Board for Plant Genetic Resources, FAO, Rome.
Gupta, V. K. and Gudu, S. (1991). Interspecific hybrids and possible phylogenetic relations in grain amaranths. Euphytica. 52: 33–38.
Gupta, A. K., Sharma, M., and Tandon N. (2004). Reviews on Indian Medicinal
Plants: Vol. 2. Indian Council of Medical Research, New Delhi, India.
Haughton, C. S. (1978). Green Immigrants: Plants that Transformed America.
Harcourt Brace Jovanovich, New York.
He, H. P. and Corke, H. (2003). Oil and Squalene in Amaranthus grain and leaf.
J Agric Food Chem. 51: 7913–7920.
He, H. P., Cai, Y. Z., Sun, M., and Corke, H. (2002). Extracion and purification
of squalene from Amaranthus grain. J Agric Food Chem. 50: 368–372.
Henderson, T. L., Johnson, B. L., and Schneiter, A. A. (2000). Row spacing,
plant population and cultivar effects on grain amaranth in the northern Great
Plains. Agronomy J. 92: 329–336.
Hoover, R., Sinnott, A. W., and Perera, C. (1998). Physiochemical characterization of starches from Amaranthus cruentus grain. Starch. 50: 456–463.
Imeri, A., Gonzalez, J. M., Flores, R., Elias, L. G., and Bressani, R. (1987). Genetic variability and correlation between yield, grain size, chemical composition and protein quality of 25 varieties of amaranth (Amaranthus caudatus).
Archivos Latinoamaricanos de Nutricion. 37: 132–146.
Irving, D. W., Betschart, A. A., and Saunders, R. M. (1981). Morphological
studies on Amaranthus curentus. J Food Sci. 46: 1170–1174.
Ivete, T. G., Egle, M. A. S., Sandra, F. A., and Elizabeth, M. T. de Souza. (2004).
Carotenoids from native Brazilian dark green vegetables and bioavailable: A
study in rats. Nutr Res. 24: 671–679.
Jain, S. K. and Defillips, R. A. (1991). Medicinal Plants of India, Vol. 1, Reference Publications Inc., Algonac, MI.
Jain, S. K. and Hauptli, H. (1980). Grain Amaranth; A new crop for California.
Agronomy progress report 107, Agric. Exp. Station, University of California,
Davis.
Jaiswal, S., Batra, A., Verma, S., and Bakodia, M. M. (1984). Free Amino acids
of some regionally available medicinally important plant seeds. Sci Cult. 50:
24–26.
Jahaniaval, F., Kakuda, Y., and Marcone, M. F. (2000). Fatty acids and triglycerol
compositions of seed oil of five Amaranthus accessions and their comparison
ton other oils. J. Am Oil Chemists Soc. 77: 847–852.
Jamriska, P. (1993). The effect of mulching with paper on seed yield of amaranth
(Amaranthus hypocondriacus L.). Rostlinna Vyroba. 39: 761–768.
Joshi, M. C., Patel, M. B., and Mehta, P. J. (1980). Some folk medicines of
Dangs, Gujarat state. Bull. Med. Ethnobot Res. 1: 8–24.
Juan, R., Pastor, J., Alaiz, M., Megias, C., and Vioque, J. (2007). Caracterizacı́on
proteica de las semillas de once especies de amaranto. Grasa Y Aceites, 58:
49–55.
Downloaded by [National Botanical Research Inst] at 04:19 22 February 2013
NUTRITIVE VALUE OF AMARANTH
Junkuszew, M., Oleszek, W., Jurzysta, M., Piancente, S., and Pizza, C. (1998).
Triterpenoid saponins from the seed of Amaranthus cruentus. Phytochemistry.
49: 195–198.
Kakrani, H. K. N. and Saluja, A. K. (1994). Traditional treatment through herbs
in Kutch district, Gujarat state, India. Part II. Analgesic, Anti-inflammatory,
antirheumatic, antiarthritic plants. Fitoterapia. 65: 427–430.
Kapoor, S. L. and Kapoor, L. D. (1980). Medicinal plant wealth of the Karimnagar district of AP. Bull Med. Ethno Bot Res. 14: 120–144.
Katiyar, R. S., Shukla, S., and Rai, S. (2000). Varietal performance of grain
amaranth (A. hypochondriacus) on sodic soil. Proc Nat Acad Sci (India). 70:
185–187.
Kauffman, C. S. and Hass, P. W. (1983). Grain Amaranth: A crop with low
water requirements and high nutritional value. In: Environmentally Sound
Agriculture, p. 299. Keretz, W. L., Ed., Praeger Press, New York.
Kong, X., Corke, H., and Bertoft, E. (2009). Fine structure characterization of
amylopectins from grain amaranth starch. Carbohydr Res. 344: 1701–1708.
Khoshoo, T. N. (1979). Cytogenetics in relation to plant evolution and Improvement. Progress in plant research. NBRI Silver Jubilee 2: 58–66.
Kumar, K. and Goel, A. K. (1998). Little known ethno-medicinal plants of
Santhal and Paharia tribals in Santhal Paragana, Bihar, India. Ethanobotany.
10: 66–69.
Kumar, A. B. S., Lakshman, B., Jayaveera, K. N., Nandeesh, R., Manoj, B.,
and Ranganayakulu, D. (2010). Comparative in vitro anthelmentic activity
of three plants from the amaranthaceae family. Arch Biol Sci, Belgrade. 62:
185–189.
Khandaker, L., Akond, A. S. M. G. M., Ali, M. B., and Oba, S. (2010). Biomass
yield and accumulations of bioactive compounds in red amaranth (Amaranthus tricolor L.) grown under different colored shade polyethylene in spring
season. Scientia Horti. 123: 289–294.
Lakshmi, B. and Vimala, V. (2000). Nutritive value of dehydrated green leafy
vegetable powders. J Food Sci Tech, Mysore. 37: 465–471.
Lee, J. R., Hong, G. Y., Dixit, A., Chung, J. W., Ma, K. H., Lee, J. H., Kang,
H. K., Cho, Y. H., Gwag, J. G., and Park, Y. L. (2008). Characterization of
microsatellite loci developed for Amaranthus hypocondiacus and their cross
amplification in wild species. Conservation Genetics. 9: 243–246.
Lehmann, J. W., Clark, R. L., and Frey, K. J. (1991). Biomass heterosis and combining ability in interspecefic and intraspecefic matings of grain amaranths.
Crop Sci. 31: 1111–1116.
Lehman, J. W. (1994). Amaranth: Commercialization and industrialization. In:
O. Paredes Lòpez (Ed.): Amaranth Biology, Chemistry and Technology, pp.
207–217. PAredes Lopez, O., Ed. CRC Press: Boca Raton FL.
Leon-Camacho, M., Garcia Gonzalez, D. L., and Aparicio, R. (2001). A detailed
and comprehensive study of Amaranth (Amaranthus curentus L.) oil fatty
profile. Eur Food Res Tech. 213:349–355.
Lindley, J. (1981). Flora Medica. Ajay Book Service, New Delhi, India.
Long, J. J., Wang, J. L., and Berry, J. O. (1994). Cloning and analysis of the C4
photosynthetic NAD dependent malic enzyme of amaranth mitochondria. J
Biological Chem. 269: 2827–2833.
Lorenz, K. (1981). Amaranthus hypochondriacus: Characteristics of the starch
and baking potential of the flour. Starch. 33: 149–153.
Lozanov, V., Guarnaccia, C., Patty, A., Foti, S., and Pongor, S. (1997). Synthesis and cystine/cysteine catalyzed oxidative folding of the Amaranth alpha
amylase inhibitor. J Peptide Res. 50: 65–72.
Legaria, J., Rajsbaum, R., Muñoz-Clares, R. A., Villegas-Sepúlveda, N., Simpson, J., and Iturriaga, G. (1998). Molecular characterization of two genes
encoding betaine aldehyde dehydrogenase from amaranth. Expression in
leaves under short-term exposure to osmotic stress or abscisic acid. Gene.
218: 69–76.
Law-Ogbomo, K. E. and Ajayi, S. O. (2009). Growth and yield performance of
Amaranthus cruentus influenced by planting density and poultry manure application. Notulae Botanicae Horti Agrobotanici Cluj-Napoca. 37: 195–199.
Madhusoodanan, K. J. and Pal, M. (1981). Cytology of vegetable Amaranths.
Bot J Linn Soc. 82: 61–68.
Mallory, M. A., Hall, R. V., McNabb, A. P., Pratt, D. B., Jellen, E. N., and
Maughan, P. J. (2008). Development and characterization of microsatellite
markers for the grain amaranths. Crop Sci. 48: 1089–1106.
123
Mathe-Gaspar, G. (2001). Green yield and drought tolerance of Amaranthus
lines. Novenytermeles. 50: 431–439.
Marcone, M. F., Niekamp, F. K., Lemaguer, M., and Yada, R. Y. (1994). Purification and characterization of the physicochemical properties of the albumin
fraction from the seeds of Amaranthus hypocondriacus. Food Chem. 51:
287–294.
Marcone, M. F., Beniac, D. R., Harauz, G., and Yada, R. Y. (1994). Quaternary structure and model for the oligomeric seed clobulin from Amaranthus
hypocondricus. J Agri Food Chem. 42: 2675–2678.
Martin, F. W. and Telek, L. (1979). Vegetables of hot humid tropics. Part 6:
Amaranth and Celosia. USDA, New Orleans, LA.
Martin, F. W. and Ruberte, R. M. (1979). Edible leaves of the tropics. U.S.
Dept. of Agriculture, Mayaguez Institute of Tropical Agriculture, Mayaguez,
Puerto Rico.
Massimo, F., Marcone, Yukiokakuda, and Yada, Y. Rikey. (2004). Amaranth a
rich dietary source of β-sitosterol and other phytosterols. Plants Food Human
Nutr. 58: 207–211.
Maughan, P. J., Sisneros, N., Luo, M. Z., Kudrna, D., Ammiraju, J. S. S., and
Wing, R. A. (2008). Construction of an Amaranthus hypocondiacus bacterial
artificial chromosome library and genomic sequences of herbicide target
genes. Crop Sci. 48: S85–S94.
McMasters, M. M., Baird, P. D., Holzapfel, M. M., and Rist, C. E. (1955).
Preparation of starch from Amaranthus cruentus seed. Econ Bot. 9: 300–302.
Medicinal Plants of Central America. www.aircav.com/survival/appb/
asappb03.html.
Medicinal Plants of Nepal (1970). HMG of Nepal, Ministry of Forests, Dept. of
Medicinal Plants, Nepal.
Michael, H. (2002). South American Medicinal Plants: Botany, Remedial Properties and General Use. Roth, I. and Lindorf, H., Eds., Springer, Berlin.
Mishra, B. (1878). Bhavprakash Nighantu. Ganga Vishnu Sri Krishna Das, Ed.,
Streem Press, Kalyan, Mumbai.
Misra, P. S., Dhan, P., Pandey, R. M., and Pal, M. (1985). Protein and amino
acids composition of grain Amaranth seeds. Fitoterpapia. 56: 318–320.
Misra, P. S., Pandey, R. M., and Pal, M. (1983). Amino acids composition in
Amaranthus. Fitoterpapia. 54:135–139.
Mohanty, R. B., Padhy, S. N., and Dash, S. K. (1996). Traditional phototherapy
for diarrhoeal diseases in Ganjan and Phulbani district of south Orissa, India.
EthnoBot. 8: 60–65.
Molina, M. I., Circosta, A., Añón, M. C., and Petruccelli, S. (2008). Mature
Amaranthus hypochondriacus seeds contain non-processed 11S precursors.
Phytochemistry. 69: 58–65.
Mukut, B. and Andhiwal, C. K. (1976). Amino acids in certain medicinal plants.
Acto Cienc. Indica. 2: 229–230.
Nadkarni, A. K. (1954). Dr. K. M. Nadkarni’s Indian Materia Medica. Vol. I
(pp. 1319) & Vol. II (pp. 968), Popular Book Depot, Bombay, India.
Nandal, S. N. and Bhatti, D. S. (1983). Preliminary screening of some weeds
shrubs for their nematicidal activity against Meloidogyne javanica. Ind J
Nemat. 13: 123–127.
NAS (1975). Underexploited Tropical Plants with Promising Economic Value.
National Academy of Science, Washington, DC.
Oke, O. L. (1983). Amaranth in Nigeria. In: Proceedings of the Second Amaranth Conference, p. 22. Rodale Press, Emmaus, PA.
Okumo, K. and Sakaguchi, S. (1981). Glutinous and non glutinous starches in
perisperm of grain amaranths. Cereal Res Commu. 9: 305.
Okumo, K., and Sakaguchi, S. (1982). Inheritance of starch characteristics in
perisperm of Amaranths hycondriacul. J Heredity. 73: 467.
Oleszek, W., Junkuszew, M., and Stochmal, A. (1999). Determination and toxicity of saponins from Amaranthus curentus seeds. J Agric Food Chem. 47:
3685–3687.
Pal, M. (1972), evolution and improvement of cultivated amaranths. I. Breeding
system and inflorescence structure. Proc Indian Nat Sci Acad. 38:28–37.
Pal, M. and Khoshoo, T. N. (1972). Evolution and Improvement of cultivated
Amaranths (v. Inviability, weakness and sterility in hybrids). J Heredity. 63:
78–82.
Pal, M. and Khoshoo, T. M. (1965). T. N. Grain amaranths. National Botanical
Gardens, Lucknow, India.
Downloaded by [National Botanical Research Inst] at 04:19 22 February 2013
124
A. RASTOGI AND S. SHUKLA
Pal, M. and Khoshoo, T. M. (1974). Grain amaranth: Evolutionary studies
in world crops: Diversity and change in Indian sub continent. Cambridge
University Press, London.
Pal, M. and Khoshoo, T. N. (1965). Origin of Amaranthus dubis. Curr Sci. 34:
370–371.
Pal, M. and Khoshoo, T. N. (1974). Grain Amaranths: Evolutionary Studies in
World Crops Diversity and Change in Indian Subcontinent. Hutchinson, J.,
Ed., Cambridge University Press, London.
Petruccelli, S., Molina, M. I., Lareu, F. J., and Circosta, A. (2007). Two
short sequences from amaranth 11S globulin are sufficient to target green
fluorescent protein and beta-glucuronidase to vacuoles in Arabidopsis cells.
Plant Physio Biochem. 45: 400–409.
Pedersen, B., Hallgren, L., Hansen, I., and Eggum, B. C. (1987). The nutritive
value of amaranth grain (Amaranthus caudatus). Plants Food Human Nutr.
36: 325–334.
Pedersen, B., Kandsen, K. E. Bach, and Eggum, B. C. (1990). The nutritive
value of amaranth grain. Plant Food Human Nutr. 40: 61–71.
Pisarikova, B., Kracmar, S., and Herzig, I. (2005). Amino acid contents and
biological value protein in various amaranth species. Czech J Animal Sci. 50:
169–174.
Prajapati, N. D., Purohit, S. S., Sharma, A. K., and Kumar, T. (2003). Handbook
of Medicinal Plants. A Canble Sourt Book, Agribios.
Prakash, D., Joshi, B. D., and Pal, M. (1995). Vitamin C in leaves and seed oil
composition of the Amaranthus species. Inter J Food Sci Nutr. 46: 47–51.
Prakash, D. and Pal, M. (1991). Nutritional and Antinutritional Composition of
vegetable and grain Amaranth leaves. J Sci Food Agric. 57: 573–583.
Pribylova, R., Pavlik, I., Rozsypalova, Z., and Bartos, M. (2006). A PCR based
for the detection of genetically modified potatoes by the gene ac2 from
Amaranthus caudatus. Eur Food Res Tech. 223: 139–142.
Pribylova, R., Kralik, P., Pisarikova, B., and Pavlik, I. (2008). Detection of the
antimicrobial peptide gene in different Amaranthus speceies. Biologia. 63:
217–220.
Praznik, W., Mundigler, N., Korler, A., Pelzl, B., and Huber, A. (1999). Molecular background of technological properties of selected starches. Starch-Strake.
51: 197–211.
Ptushenko, V. V., Gins, V. K., and Tikhonov, A. N. (2002). Interaction of amaranthin with the elect transport chain of chloroplasts. Russian Plant Physiology.
49: 585–591.
Pandey, R. M. (2009). Genetic divergence of parents and F-2 segregation in
grain Amaranths. Cientia E Investigacion Agrari. 6: 77–84.
Palado, M. C. and Chang, L. C. (2003). Suggested cultural practices for vegetable amaranth. AVRDG. Publication No. 03-552.
Park, Y. J., Nemoto, K., Nishikawa, T., Matsushima, K., Minami, M., and
Kawase, M. (2009). Molecular cloning and characterization of granule bound
starch synthase I cDNA from a grain amaranth (Amaranthus cruentus L.).
Breed Sci. 59: 351–360.
Qian, J. Y. and Kuhn, M. (1999). Characterization of Amaranthus cruentus and
Chenopodium quinoa starch. Strach Strake. 51: 116–120.
Radek, M. and Savage, G. P. (2008). Oxalates in some Indian green leafy
vegetables. Inter J Food Sci Nutr. 59: 246–260.
Raina, A. and Dutta, A. (1992). Molecular cloning of a gene encoding a seed
specific protein with nutrionally balanced amino acid composition from Amaranthus. Proceedings National Academy Sci, USA. 89: 11774–11778.
Raj, K. P. S. and Patel, M. R. (1978). Some medicinal plants of cambay and its
immediate vicinity and their uses in Indian Indigenous system of medicine.
Ind Drugs. 15: 145–152.
Rodas, B. and Bressani, R. (2009). The oil, fatty acid and squalene content of
varieties of raw and processed grain amaranth. Archivos Latinoamericanos
De Nutricion. 59: 82–87.
Rajwar, G. S. (1983). LowAltitude medicinal plants of south Garhwal. Bull Med
Ethnobot. Res. 4: 14–28.
Rodriguez, S. V., Segura-Nieto, M., Chagolla-Lopez, A., Vargas-Cortina, A. V.,
Martinez-Gallardo, N., and Blanco-Labra, A. (1993). Purification, characterization, and complete amino acid sequence of a trypsin inhibitor from
Amaranth (Amaranthus hypochondriacus) seeds. Plant Physiology. 103:
1407–1412.
Rodrı́guez, S.V., Guerrero-Rangel, A., Melgoza-Villagómez, C., ChagollaLópez, A., Delgado-Vargas, F., Martı́nez-Gallardo, N., Sánchez-Hernández,
C., and Délano-Frier, J. (2007). Cloning of a cDNA encoding a cystatin from
grain amaranth (Amaranthus hypochondriacus) showing a tissue-specific expression that is modified by germination and abiotic stress. Plant Physio
Biochem. 45: 790–798.
Ramachandran, M. and Phanasalkar, P. V. (1956). Essential amino acid composition of certain vegetable food stuffs. Indian J Med Res. 44: 501–509.
Ramirez-Medeles, M. D., Aguilar, M. B., Miguel, R. N., BolanosGarcia, V. M.,
Garcia Hernandez, E., and Soriano Garcia, M. (2003). Amino acid sequence,
biochemical characterization and comparative modelling of a nonspecific
lipid transfer protein from Amaranthus hypocondriacus. Archives Biochem
Biophy. 415: 24–33.
Raina, A. and Datta, A. (1992). Molecular cloning of a gene encoding a seed
specific protein with nutritionally balanced amino acid composition from
amranthus. Proc Natl Acad Sci, USA. 89: 11774–11778.
Ranade, S. A., Kumar, A., Goswami, M., Farooqui, N., and Sane, P. V. (1997).
Genome analysis of amaranths: Detection of inter and intra species variations.
J Biosci. 22: 457–464.
Rangarajan, A., Chenoweth, W. A., Kelly, J. F., and Agee, K. M. (1998). Iron
bioavailability from Amaranthus species: 2-Evaluation using haemoglobin
repletion in anaemic rats. J Sci Food Agric. 78: 274–280.
Ray, T. and Roy, S. C. (2007). Phylogenetic relationships between members
of amaranthaceae and chenopodiaceae of lower gangetic plains using RAPD
and ISSR markers. Bangaladesh J Bot. 36: 21–28.
Reddy, V. V. P. and Varalakshmi, B. (1998). Heterosis and combining ability for
leaf yield and its components in vegetable amaranth (Amaranthus tricolor).
Ind J Agric Sci. 68: 773–775.
Reddy, M. B., Reddy, K. R., and Reddy, M. N. (1989). A survey of plant crude
drugs of Anantapur district, A.P., India. Int J Crude Drug Res. 27:2417–2420.
Repo-Carrasco-Valencia, R., Hellstrom, J. K., Pihlava, J. M., and Mattila P.
H. (2010). Flavonoids and other phenolic compounds in Andean indigenous
grains: Quinoa (Chenopodium quinoa), kaniwa (Chenopodium pallidicaule)
and kiwicha (Amaranthus caudatus). Food Chem. 120: 128–133.
Resio, A. C., Agurre, R. J., and Saurez, C. (1999). Analysis of the sorptional
characteristics of amaranth starch. J Food Engi. 42: 51–57.
Robert, Y. N., Hiroe, K., and Yotaro, K. (2008). Antioxidant activity of various
extracts and fractions of Chenopodium quinoa and Amaranthus spp. seeds:
Analytical, nutritional and clinical methods. Food Chem. 106: 760–766.
RomeroZepeda, H. and ParedesLopez, O. (1996). Isolation and characterization of amarantin the 11s amaranth seed globulin. J Food Biochem. 19:
329–339.
Roy, S., Dutta, A. K., and Chakraborty, D. P. (1982). Amasterol an ecdysome
precursor and a growth inhibitor from Amaranth viridis. PhytoChem. 21:
2417–2420.
Sankarnarayanan, A. S. (1988). Folk-lore medicines for jaundice from Coimbatore and Palghat district of Tamil Nadu and Kerala, India. Ancient Sci Life.
7: 175–179.
Sarkar, R., Nandan, C. K., Mandal, S., Patra, P., Das, D. I., and Syed, S. (2009).
Structural characterization of a heteropolysaccharide isolated from hot water
extract of the stems of Amaranthus tricolor Linn. (Amaranthus gangeticus
L.). Carbohydr Res. 344: 2412–2416.
Sauer, J. D. (1967). The grain Amaranths and their relatives: A revised taxonomic and geographic survey. Ann. Missouri Bot Gard. 54: 103–137.
Sauer, J. D. (1979). Grain Amaranths. In: Evolution of Crop Plants, pp. 4–7.
Simmonds, N.W., Ed., Longman, London.
Sauer, J. D. (1950a). Amaranths as dye plants among the pueblo peoples. Southwest J Anthropol. 6: 412–415.
Sauer, J. D. (1977). The history of grain Amaranths and their uses and cultivation
around the world. In: Proceedings of the first Amaranth Seminar, p. 9. Rodale
Press, Emmaus, PA.
Sauer, J. D. (1950b). The grain amaranths: A survey of their history and classification. Ann Missouri Bot Gard. 37: 561–632.
Saunders, R. M. and Becker, R. (1984). Amaranthus; A potential food and feed
resource. In: Advanced Cereal Science Technology, Vol. 6, p. 357, Pomeranz,
Y., Ed. American Association of Cereal Chemists, St. Paul, MN.
Downloaded by [National Botanical Research Inst] at 04:19 22 February 2013
NUTRITIVE VALUE OF AMARANTH
Sebasttian, M. K. and Bhandari, M. M. (1984). Medicoethno botany of Mount
Abu, Rajasthan, India. J Ethanopharmacol. 12: 223–230.
Senft, J. P. (1980). Protein quality of Amaranth grain. In: Proceedings of the
Second Amaranth Conference, p. 43. Rodale Press Inc., Emmaus, PA.
Sharma, J. K., Lata, S., and Sharma, R. P. (2001). Stability for grain yield in
amaranth (Amaranthus hypocondriacus). Ind J Agric Sci. 71: 392–394.
Siddiqui, M. A. and Alam, M. M. (1997). Organic soil amendments withsome
non-conventional plant additives for the management of Meloidogyne incognita infecting tomato. Ind J Nemato. 27: 120–122.
Siddiqui, M. B., Alam, M. M., and Husain, W. (1989). Traditional treatment of
skin diseases in Uttar Pradesh, India. Econ Bot. 43: 480–486.
Siener, R., Honow, R., Seidler, A., Voss, S., and Hesse, A. (2006). Oxalate contents of species of the Polygonaceae, Amaranthaceae and Chenopodiaceae.
Food Chem. 98: 220–224.
Singh, B. P. and Whitehead, W. F. (1996). Management methods for producing
vegetable amaranth. In: Progress in New Crops, pp. 511–515. Janick, J., Ed.,
ASHS Press, Arlington, VA.
Singh, V. K. and Dhar, U. (1993). Folk medicines of Orissa: Keonjhar forests.
Glimpse Plant Res. 10: 103–107.
Singh, S. K., Das, P. K., and Das, M. N. (1998). Ethno-medical studies on some
medicinal plantsn of Rajgir, Bihar. Int J Mendel. 15: 145–148.
Sharma, R. K. and Mukut, B. (1991). Screening of the compounds isolated
from Amaranthus tricolor for antibacterial activity. Acta Ciene Indica. 17:
357–362.
Shukla, S. and Singh, S. P. (2003). Correlation and path analysis in grain amaranth (Amaranthus spp.). Ind J Genetics. 63: 163–164.
Shukla, S., Bhargava, A., Chatterjee, A., Pandey, A. C., and Mishra, B. K.
(2010). Diversity in phenotypic and nutritional traits in vegetable amaranth
(Amaranthus tricolor): A nutritionally underutilized crop. J Sci Food Agric.
90: 139–144.
Shukla, S., Bhargava, A., Chatterjee, A., Srivastava, A., and Singh, S. P. (2005).
Estimates of genetic variability in vegetable amaranth (A. tricolor) over different cuttings. Horti Sci (Prague). 32: 60–67.
Shukla, S. and Singh, S. P. (2001). Correlated responses in amaranth. Ind J Plant
Genetic Resources. 14: 371–372.
Shukla, S. and Singh, S. P. (2002). Genetic divergence in amaranth (Amaranthus
hypocondiacus L.). Ind J Genet. 62: 336–337.
Shukla, S., Pandey, V., Pachauri, G., Dixit, B. S., Banerji, R., and Singh, S. P.
(2003). Nutritional contents of different foliage cuttings of vegetable amaranth. Plant Food Hum Nutr. 58: 1–8.
Shukla, S., Bhargava, A., Chatterjee, A., and Singh, S. P. (2004). Estimates of
genetic parameters to determine variability for foliage yield and its different
quantitative and qualitative traits in vegetable amaranth (A. tricolor). J Genet
Breed. 58: 169–176.
Shukla, S., Bhargava, A., Chatterjee, A., Srivastava, A., and Singh, S. P. (2006).
Genotypic variability in vegetable amaranth (Amaranthus tricolor L.) for
foliage yield and its contributing traits over successive cuttings and years.
Euphytica. 151: 103–110.
Shukla, S. and Singh, S. P. (2000). Studies on genetic parameters in vegetable
amaranth. J Genet Breed. 54: 133–135.
Shukla, S., Bhargava, A., Chatterjee, A., Srivastava, J., Singh, N., and Singh, S.
P. (2006). Mineral profile and variability in vegetable Amaranth (Amaranthus
tricolor). Plant Foods Human Nutr. 61: 23–28.
Srivastava, T. N., Rajasekharan, S., Badola, D. P., and Shah, D. C. (1986). An
index of the available medicinal plants used in Indian system of medicine
from Jammu and Kashmir State. Ancient Sci Life. 6: 49–63.
Stallknecht, G. F., and Schulz-Schaeffer, J. R. (1993). Amaranth rediscovered.
In: Janick J. and Simon, J.E. (eds.), pp. 211–218, New Crops. New YorkWiley.
Stefunova, V. and Bezo, M. (2003). Genetic diversity analysis of amaranth
(Amaranth Cruentus) germplasm collection by RAPD. Biologia. 58:53–57.
Stintzing, F. C., Kammerer, D., Schieber, A., Adama, H., Nacoulma, O. G., and
Carle, R. (2004). Betacyanins and phenolic compounds from Amaranthus
spinosus L. and Boerhavia erecta L. J Biosci. 59: 1–8.
Stone, L. A. and Lorenz, K., (1984). The starch of Amaranthus: Physiochemical
properties and functional characteristics. Starch. 36: 232–237.
125
Sudhakar, A. and Chetty, K. M. (1998). Medicinal importanceof some angiosperm weeds used by the rural people of Chittor district of A.P., India.
Fitoterpia. 69: 390–400.
Sumar, L. S. (1983). The small giant. Amaranth Newsletter, Archieves Latino
Americanos de Nutricion. 2: 1.
Suma, S., Ambika, S. R., Kazinczi, G., and Narwal, S. S. (2002). Allelopathic
plants. 6. Amaranth spp. Allelopathy J. 10: 1–11.
Sun, M., Chen, H., and Leung, F. C. (1999). Low cot DNA sequences for fingerprinting analysis of germplasm diversity and relationships in Amaranthus.
Theoretical and Applied Genetics. 99: 464–472.
Suresh, B., Dhanasenkaran, S., Kumar, R. V., and Balasubramanian, S. (1995).
Ethanopharmacological studies on the medicinal plants of Nilgiris Ind Drugs.
32: 340–352.
Tomoskozi, S., Baracskai, I., Schonlechner, R., Berghofer, E., and Lasztity,
R. (2009). Comparative study of composition and technological quality of
amaranth I. Gross chemical composition, amino acid and mineral content.
Acta Alimentaria. 38: 341–347.
Tamir, S., Bell, J., Finlay, T. H., Sakal, E., Smirnoff, P., Gaur, S., and Birk, Y.
(1996). Isolation, characterization, and properties of a trypsin-chymotrypsin
inhibitor from amaranth seeds. J Protein Chem. 15: 219–229.
Teutonico, R. A. and Knorr, D. (1984a). Plant tissue culture: Food applications and the potential reduction of nutritional stress factors. Food Tech. 38:
120–123.
Teutonico, R. A. and Knorr, D. (1984b). Food Potential of Amaranth Tissue
Culture. Plant Cell Reports.
Teutonico, R. A. and Knorr, D. (1985). Non destructive method for oxalate
determination of cultured Amaranthus tricolor cells. J Agric Food Chem. 33:
60–64.
Thellung, A. (1914). Amaranthus. In: F. Ascherson and P. Graebner, Synopsis
der Mittleuropäischen Flora, Vol. 5, pp. 225–356. Ascherson, F. and Graebner, P., Eds. Borntraeger, Leipzig (Engelmann).
Tomita, Y., Sugimoto, Y., Sakomoto, S., and Fuwa, H. (1981). Some properties
of starches of grain Amaranthus ans several millets. J Nutr Sci Vitaminol. 27:
471–484.
Transue, D. K., Fairbanks, D. J., Robinson, L. R., and Andersen, W. R. (1994).
Species identification by RAPD analysis of grain amaranth genetic resources.
Crop Sci. 34: 1385–1389.
Tripathi, Y. C., Prabhu, V. V., Pal, R. S., and Mishra, R. N. (1996). Medicinal
plants of Rajasthan in India system of medicine.Ancient Sci Life. 15: 190–
212.
Vaidyaratnam, P. S. V. (1993). Indian Medicinal Plants, Vol. 1, Arya Vaidya
Sala.
Valdés-Rodrı́guez, S., Blanco-Labra, A., Gutiérrez-Benico, G., Boradenenko,
A., Herrera-Estrella, A., and Simpson, J. (1999). Cloning and characterization
of a trypsin inhibitor cDNA from amaranth (Amaranthus hypochondriacus)
seeds. Plant Mol. Biol. 41: 15–23.
Varalakshmi, B. (2003). Phenotypic stability for economic traits in vegetable
amaranth (Amaranth tricolor). Ind J Agric Sci. 73: 114–115.
Vasi, I. G. and Kalintha, V. P. (1980). Amino acids composition of some leafy
vegetables. J Inst Chem, India. 52: 13–14.
Vietmeyer, N. (1978). Poor people’s crops. Agenda AID. 1: 12–17.
Wetzel, D. K., Horak, M. J., and Skinner, D. Z. (1999). Use of PCR based
molecular markers to identify weedy Amaranthus species. Weed Sci. 47:
518–523.
Wu, H. X., Sun, M., Yue, S. X., Sun, H. L., Cai, Y., Huang, R. H., Brenner, D.,
and Corke, H. (2000). Field evaluation of an Amaranthus genetic resource
collection in China. Gene Resou Crop Evol. 47: 43–53.
Wu, H. X. and Corke, H. (1999). Genetic diversity in physical properties of
starch from a world collection of Amaranthus. Cereal Chem. 76: 877–883.
Xu, F. X. and Sun, M. (2001). Comparative analysis of phylogenetic relationships of grain Amaranthus and their wild relatives (Amaranthus; amaranthaceae) using internal transcribed spacer, AFLP and double primer florescent
intersimple sequence repeat markers. Mole Phylogenetics Evol. 21: 372–387.
Yoganarasimthan, S. N. (2000). Medicinal Plants of Tamil Nadu, Vol. 2.
www.eap.mcgill.ca/CPAT 1.htm. Nov. 2008. www.wikepedia.com. USDA National Research Council.