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Cultivated Ancient Wheats ( Triticum spp.): A Potential Source of Health-Beneficial Food Products: Ancient wheats for healthy foods…

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A sustainable and wholesome food supply is the most important incentive that has led to an increasing interest in ancient wheats over the past few decades. Domestication of wheat, followed by breeding efforts, largely over the past 2 centuries, has resulted in yield increases but with grain quality deterioration due to the reduction of protein, vitamins, and minerals in grains. It has also resulted in a decrease in food diversity due to the loss of genetic variation in the cultivated wheat gene pool. Ancient hulled wheats, einkorn, emmer, and spelt are among the early cereals that were domesticated in their places of origin in the Fertile Crescent of the Middle East where their wild predecessors still grow. The ancient wheats had a long history as part of human diet, and played an important role as a major source of food for the early civilizations in that region. The risks of genetic erosion of crop plants and the associated likely consequences for agriculture now call for revitalization of the unrealized potentials of ancestral species like einkorn, emmer, and spelt wheat, the domesticated ancestors of modern durum and bread wheats. These ancestors need to be exploited to maximize the sustainable supply of grain protein, fiber, minerals, and phytochemicals. In addition, ancient wheat biodiversity can be utilized to ensure sustainable wheat production in the context of climate change and low-input organic farming systems. This review provides a holistic synthesis of the information on ancient wheats to facilitate a greater exploitation of their potential benefits.
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Cultivated Ancient Wheats (Triticum spp.):
A Potential Source of Health-Beneficial
Food Products
Ahmad Arzani and Muhammad Ashraf
Abstract: A sustainable and wholesome food supply is the most important incentive that has led to an increasing
interest in ancient wheats over the past few decades. Domestication of wheat, followed by breeding efforts, largely over
the past 2 centuries, has resulted in yield increases but with grain quality deterioration due to the reduction of protein,
vitamins, and minerals in grains. It has also resulted in a decrease in food diversity due to the loss of genetic variation in
the cultivated wheat gene pool. Ancient hulled wheats, einkorn, emmer, and spelt are among the early cereals that were
domesticated in their places of origin in the Fertile Crescent of the Middle East where their wild predecessors still grow.
The ancient wheats had a long history as part of human diet, and played an important role as a major source of food for
the early civilizations in that region. The risks of genetic erosion of crop plants and the associated likely consequences
for agriculture now call for revitalization of the unrealized potentials of ancestral species like einkorn, emmer, and spelt
wheat, the domesticated ancestors of modern durum and bread wheats. These ancestors need to be exploited to maximize
the sustainable supply of grain protein, fiber, minerals, and phytochemicals. In addition, ancient wheat biodiversity can be
utilized to ensure sustainable wheat production in the context of climate change and low-input organic farming systems.
This review provides a holistic synthesis of the information on ancient wheats to facilitate a greater exploitation of their
potential benefits.
Keywords: diversity, einkorn, emmer, flour, grain, nutrient rich food, spelt, Triticum spp
Introduction
Common wheat, bread and soft wheat (Triticum aestivum L. em
Thell; AABBDD; 2n=6x=42), was the world’s most important
crop in 2014 with a world production of about 730 million tons
harvested from an area of over 220 million ha (FAOSTAT 2016).
Used as a major staple food, as bread, in many countries, it has been
the most abundant source of calories and protein in the human
diet, supplying nearly 20% of the total dietary protein worldwide
(Braun and others 2010). Durum or macaroni wheat (Triticum
turgidum L. subsp. durum Desf.) is tetraploid wheat (AABB;
2n=4x=28) which is widely used for the production of pasta.
Common wheat is the most widely grown species accounting
for 95% of the total with durum wheat representing the remain-
ing 5%. Hulled wheats with nonfragile spikes and hulled grains
were the earliest species domesticated, almost 10000 y ago during
the Neolithic period. These played a key role in the phylogen-
esis of modern wheat. Easy cultivation, harvestability, and long-
term storage capacity of wheat grains enabled man to establish
CRF3-2017-0018 Submitted 1/24/2017, Accepted 2/25/2017. Author Arzani
is with Dept. of Agronomy and Plant Breeding, College of Agriculture, Isfahan Univ.
of Technology, Isfahan 84156-83111, Iran. Author Ashraf is with Pakistan Sci-
ence Foundation, Islamabad, Pakistan. Direct inquiries to author Arzani (E-mail:
a_arzani@cc.iut.ac.ir).
his early settlements at the dawn of civilization as populations
increased in the Babylonian and Assyrian empires over the area
that encompasses the “Fertile Crescent.” Wheat as a staple food
was cultivated in ancient Persia, Greece, and Egypt in prehistoric
times.
The global distribution and utilization of wheat is a con-
sequence of its unique dough rheological properties and the
bread-baking quality of its flour. A risen dough can be baked
into a loaf of bread because of the characteristic physicochemical
properties of gluten that provides the dough with the required
elasticity. The gluten network can be stretched as a result of
CO2gas bubbles trapped inside the fermenting dough, which is
critical in the production of a variety of breads and fermented
products. Pasta quality also largely depends on the gluten protein
composition and content in durum wheat grains.
Ancient cultivated wheats (einkorn, emmer, and spelt) origi-
nated in the Fertile Crescent, an area in the Middle East spreading
from Jordan, Palestine, and Lebanon to Syria, Turkey, Iraq, and
Iran (Figure 1), where their wild ancestors are still found (Harlan
and Zohary 1966). The ancient wheats have been recognized as
a primary component of the human diet in the Old World dur-
ing the Bronze and Neolithic ages. They represented a strategically
important food crop among the ancient Assyrian, Babylonian, and
Egyptian nations (Figure 1). These hulled (glumed) wheats com-
prise all 3 polyploidy levels of diploid (2x), tetraploid (4x), and
C2017 Institute of Food Technologists®
doi: 10.1111/1541-4337.12262 Vol.16, 2017 rComprehensiveReviews in Food Scienceand Food Safety 477
Ancient wheats for healthy foods . . .
Figure 1–This is the region where some of the most ancient agricultural communities in the world came into being in the Fertile Crescent. The
domestication of wheat and barley occurred thousands of years ago in this region. It also comprises Sumer, the birthplace of the 1st civilization and
the Bronze Age; and Mesopotamia consisting of Sumerian, Babylonian, Akkadian, and Assyrian empires (Adapted from:
http://resourcesforhistoryteachers.wikispaces.com/7.7).
hexaploid (6x) present in Triticum spp. Einkorn (T. monococcum L.)
is a diploid wheat (AA; 2n=2x=14) which is now cultivated in
limited regions in the world (Arzani 2011). Emmer (T. turgidum L.
spp. dicoccum Schrank ex Sch¨
ubler) is a tetraploid wheat (AABB;
2n=4x=28) which is a domesticated form of T. turgidum spp.
dicoccoides (wild emmer wheat). Triticum turgidum ssp. durum (Desf)
Husn. (durum wheat) originated from the domesticated emmer
(Figure 2). Spelt (T.aestivum subsp. spelta) is a hexaploid wheat
(AABBDD; 2n=6x=42) and is most likely the ancestor of the
free-threshing common wheat. Thus, einkorn, emmer, and spelt
represent the 3 cultivated species of hulled wheat which include a
bridging species between the cultivated (bread wheat and durum
wheat) and wild wheats (Figure 2). Einkorn, emmer, and spelt
wheat types are the earliest cultivated ones; hence, designated as
ancient wheats.
Over the centuries, desirable bread quality was achieved through
a variety of traditional methods effectively using available raw
materials. In some places, the traditional craft of bread-making
has been conserved, while in others it has undergone substantial
change. For example, steamed breads in China and flat breads in
the Middle East are still traditionally baked in large quantities up-
holding ancestral cultural traditions. North America, in contrast,
has witnessed the development of new wheat cultivars since its
introduction by European settlers, while innovative developments
introduced by the bread-making industry have replaced maize as
the traditional staple crop.
The whole wheat grain is milled and usually only the endosperm
is used to make white flour, while the bran and germ are removed
as by-products (see review by Hemdane and others 2016). Para-
doxically, it is the whole grain that is rich in protein, minerals, and
vitamins, with the refined grain largely consisting of starch. Both
the common and durum wheats have the free-threshing forms
in which the glumes are fragile and the rachis tough. On the
other hand, the grains of the 3 ancient domesticated species of
wheat (einkorn, emmer, and spelt) are covered by glumes (hulls)
even after harvest. The disadvantage with these ancient cultivated
wheats is that their grains are difficult to thresh because the hulls
remained attached upon threshing so that further processing is
required to remove the hulls or husks and make them ready for
milling or pounding. The toughened glumes provide protection
against pests of stored grain since these types of wheat are usually
stored as spikelets.
Nutritional Qualities of Modern and Ancient Wheats
Evidence shows that the proportion of the global human pop-
ulation facing malnutrition due to inadequate micronutrients has
risen over the past several decades (Miller and Welch 2013; Kumar
and others 2016). This is believed to be partly the consequence
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Ancient wheats for healthy foods . . .
Figure 2–Phylogeny of domesticated species of the Triticum spp., including einkorn, emmer, spelt, durum, and common wheat.
of replacement of not only “ancient” but also “old” crop cul-
tivars. It is well known that modern and old common wheats
(T.aestivum) differ significantly from each other in terms of their
micronutrients, with zinc and iron specifically being lower in mod-
ern cultivars (Fan and others 2008). The most commonly quoted
evidence supporting this claim comes from a long-term study, of
more than 160 y, conducted at Rothamsted Research (Harpenden,
U.K.), which is further confirmed by several other studies evalu-
ating old and modern wheat cultivars/lines in modern field tri-
als. Although some evidence indicating the superiority of the
old wheat over the modern one in terms of other grain quality
components is scant, it is believed that intensive breeding might
have led to lower concentrations of minerals in modern wheat
grains due to the “yield dilution phenomenon” (Shewry and oth-
ers 2016). Therefore, in the light of current evidence, there is
no doubt that old common wheats are superior to modern ones
due to their higher micronutrient contents (Garvin and others
2006; Fan and others 2008; Shewry and others 2016). Also, in-
creased starch content may lead to the reduction of other grain
components in modern wheat cultivars. This is further supported
by the fact that increased grain yield is clearly associated with a
rise in harvest index as a result of employing modern short-straw
cultivars. The semi-dwarf wheat phenotype in common wheat
(T.aestivum L.) was developed by the introduction of functionally
constitutive growth repressor (Rht) genes which resulted in en-
hanced efficiency of partitioning photosynthetic assimilates into
grain.
An alternative or supplementary hypothesis would be the smaller
root system due to the dwarfism that may have negatively influ-
enced the capacity of the plant to absorb micronutrients from
the soil, or its capacity to store them in the vegetative organs
before being remobilized to the grain. Nonetheless, the impor-
tance and improved health benefits of wheat in the dietary chain
of many people in the world imply that wheat breeders should
target the concentration of beneficial ingredients in the grain
of novel cultivars ideally using ancient wheat species. However,
both ancient and old wheats need to be exploited to achieve bio-
fortification aimed at enhancement of micronutrient contents and
their bioavailability in grains.
Malnutrition, or nutritional deficiency, is one of the main rea-
sons underlying the high morbidity and mortality rates especially
in poor households that, compared with those with higher in-
comes, are more likely to eat more starch than protein from meat
or other sources. The need for crop diversification, the increasing
demand for nutritionally healthy food products, and the asserted
therapeutic properties of foodstuff have led to a renewed interest
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Ancient wheats for healthy foods . . .
in ancient wheats such as einkorn (T. monococcum L.), emmer (T.
turgidum ssp. dicoccum), or spelt (T. spelta). The global increasing
incidences of obesity and metabolic diseases related to diet have
resulted in a great interest in the ancient food diet. Recent evo-
lutionary diet studies have suggested that our present-day diets
should somehow bear resemblance to those of the Palaeolithic hu-
man diets containing carbohydrate-rich food derived from both
underground starchy tubers like potatoes and starchy grains such
as wheat, barley, oats, rye, millet, and maize (Hardy and others
2015; Solon-Biet and others 2016).
The common wheat most likely originated from spelt wheat
through hybridization between the emmer wheat and the diploid
goatgrass (Aegilops tauschii, DD genome) (Figure 2; Arzani and
others 2005). However, domestication of wheat clearly occurred
as a result of selecting 2 traits of reduced shattering of the spike at
maturity and free-threshing. While grain shattering is undoubtedly
an essential trait of the wild species responsible for the effective
dispersion of seeds, nonshattering through a “tough rachis” is
governed by the br (brittle rachis) gene in wheat (Nalam and
others 2006). Governed by a recessive gene tg (tenacious glume),
a modifier dominant gene Qand several other mutations in other
loci (Dvorak and others 2012), the free-threshing trait of wheat
due to the loss of strong glumes is one of the most important
factors contributing to the domestication of wheat. This trait not
only helps discriminate between “spelt and common” or between
“emmer and durum” wheats but also facilitates the management,
harvest, and use of the grains.
Wheat grains mostly contain carbohydrates, proteins, lipids, and
minerals (Table 1). In what follows, each of these ingredients is
explored.
Proteins
Wheat proteins can be classified into the 2 major gluten
(glutenin and gliadin) and nongluten (globulin and albumin) frac-
tions. The polymeric glutenins have the ability to impart strength
and elasticity to dough, whereas the gliadins are monomeric
proteins responsible for dough viscosity (D’Ovidio and Masci
2004). Based on electrophoretic mobility at low pH, glutenins are
further subdivided into low-molecular-weight (LMW) and high-
molecular-weight (HMW) proteins. According to the repetitive
amino acid sequence patterns, gliadins are classified into the 4 types
of α-, β-, γ-, and ω- (Barak and others 2015). When subjected
to gastrointestinal digestion, each component of wheat grain
protein breaks down into a vast variety of peptides with variable
lengths. Since gluten is rich in proline, compact and tough
structures are generated that commonly pose difficulties in
digestion and elimination (Arentz-Hansen and others 2000).
Some of these digestion-resistant peptides are believed to be
involved in unfavorable immune processes in susceptible persons.
For example, one of the key immunotoxic peptides for celiac
patients is gliadin 33-mer peptide of gluten which is resistant to
digestion (Shan and others 2002).
Grain quality attributes, including its protein content, not only
vary within and across species but are strongly influenced by the
environment as well (Ashraf 2014; Arzani and Ashraf 2016). Com-
parisons of ancient and common wheats in Table 1 reveal that grain
protein contents (whole grain flour) of the ancient wheat are com-
monly superior to those of the modern counterpart (Abdel-Aal
and others 1995; Ranhotra and others 1996; Loje and others
2003; Marconi and Cubadda 2005; Brandolini and others 2008;
Shewry and others 2013). Grain size and weight are important
determinants in many qualitative and compositional constituents
such that a big and heavy grain, for instance, yields a larger en-
dosperm as well as smaller proportions of aleuronic layers and
external pericarp. A wheat grain includes 3 main components of
germ (embryo), endosperm, and outer layers (pericarp, seed coat
and, aleurone). The endosperm accounts for approximately 90%
of the dry weight of the grain. Compared to the ancient wheat,
the lower protein content of modern wheat may be explained by
its bigger and heavier grain that yields a larger starchy endosperm
which, in turn, lessens its protein content.
Carbohydrates
Carbohydrates account for the most abundant fractions (about
65% to 75%) of the wheat grain, with starch being the main con-
stituent of wheat grain and flour (approximately 60% to 70%)
(Lineback and Rasper 1988). Starch is composed of amylose
and amylopectin which are α-D-glucose polymers of different
structures intertwined to form a starch granule. Amylose is eas-
ily hydrolyzed by amylase enzymes (α-amylase and β-amylase) to
maltose, while amylopectin is degraded to maltose (approximately
60%) and dextrins (approximately 40%). Despite the fact that amy-
lose accounts for only around one-quarter of the starch granule,
the frequency of its molecules is almost 150 times greater than
that of amylopectin ones. This is due to the much smaller size of
amylose compared to amylopectin. On the other hand, its higher
resistance to hydrolytic enzymes, due to the more tightly packed
amylose chains than amylopectin ones, leads to the low postpran-
dial levels of glycemic and insulinemic responses which, in turn,
give rise to reduced postmeal blood insulin and glucose, yielding
longer satiety.
A number of studies (including the present one as reported
in Table 1) show either a comparable or even lower starch
content of ancient wheat compared with that of bread wheat
(Mohammadkhani and others 1998; Rodriguez-Quijano 2004;
Brandolini and others 2008; Caballero and others 2008;
Haghayegh and Schoenlechner 2010). The enhanced grain
yield of modern wheat results in the so-called “yield dilution
phenomenon” coupled with the higher ploidy level may explain
the higher starch content of modern wheat.
Most of the starches from white bread and white rice are read-
ily digested in the small intestine, leading to a fast rise in blood
glucose, which may be linked to the risk of developing obesity.
Type-2 diabetes has been documented in people who frequently
eat starch-based diets such as the Asian ones (Hu and others 2012).
In addition, ample evidence from both human and animal inves-
tigations support a link between wheat gluten and type-1 diabetes
(for a review, see Barbeau 2012). However, a fraction of starch,
namely, the resistant starch (RS), resists digestion or breakdown
by α-amylase in the stomach and may be delivered from the small
intestine to the colon where it undergoes fermentation to yield
small-sized fatty acids, such as butyrate, which may be beneficial
to health because it can lower colorectal cancer risks (Topping
2007).
In 2011, global human consumption of wheat was estimated to
be 65 kg per head per annum, with the highest estimate of 210 kg
observed in Azerbaijan (Bilgic and others 2016). Considering the
fact that wheat is the primary staple food for over one-third of
the world population, the reintroduction of ancient wheat species
can be pondered for a variety of reasons including the nutritional
enrichment and diversification of food. A plethora of literature
sources has been published in recent years expounding the em-
ployment of genetic resources in plant breeding. Recently, Arzani
and Ashraf (2016) addressed the need for the sustainable use of
480 ComprehensiveReviews in Food Scienceand Food Safety rVol. 16, 2017 C2017 Institute of Food Technologists®
Ancient wheats for healthy foods . . .
Table 1–Mean or range of macronutrients of whole-grain flour in cultivated einkorn, emmer, spelt, and bread wheats.
Component (dry matter) Einkorn Reference@Emmer Reference Spelt Reference Bread wheat Reference
Digestible carbohydrate (% or g per 100 g) 64.5 (1) 71 (2) 65.9(1) 73(3)
Starch (%) 62.3(1; 4; 5) 65 (2) 63.8(1) 68.5 (4)
Amylose (% starch) 23.8(4; 6) 25.1 (6) () 28.4(7)
Dietary fiber (%) 9.8(8; 9) 9.8 (9) 12 (9) 13.4 (10)
Protein (%) 15.5–22.8 (4; 11) 13.5–19.05 (8; 11) 16.3–17.5 (1) 12.9 –19.9 (12)
Lipid (%) 3.5(13; 14) 2.16(11; 14) 2.39(1; 14) 2.8 (13)
Ash (%) 2.3 (8; 13) 2.3 (8) 2.1 (8) 1.9 (8)
Phosphorus (g kg1) 5.2 (15) 5.12 (15) 4.70 (15) 4.18 (15)
Potassium (g kg1) 4.29 (15) 4.39 (15) 4.17 (15) 5 (15)
Sulfur (g kg1) 1.90 (1; 15) 1.88 (15) 1.8 (15) 1.4 (15)
Magnesium (g kg1) 1.63 (15) 1.67 (15) 1.5 (15) 1.44 (15)
Calcium (g kg1) 0.42 (15) 0.36 (15) 0.39 (15) 0.43 (15)
Iron (mg kg1) 49 (15) 49 (15) 50 (15) 38 (15; 16)
Zinc (mg kg1) 53 (15) 54 (15) 47 (15) 35.0 (15)
Manganese (mg kg1) 28 (15) 24 (15) 27 (15) 26 (15)
Copper (mg kg1) 4 (15) 4.1 (15) 5 (15) 3.9 (15)
Sodium (mg kg1) 7 (15) 12 (15) 10 (15) 10 (15)
By taking an average value either from the genotypes reported in a reference or taking an average from the listed references.
@References: 1. Abdel-Aal and others (1995); 2. Lacko-Bartosova and Curna (2015); 3. Davis and others (1981); 4. Brandolini and others (2008); 5. Brandolini and others (2011); 6. Mohammadkhani and others
(1998); 7. Regina and others (2015); 8. Loje and others (2003); 9. Gebruers and others (2008); 10. Andersson and others (2013); 11. Grausgruber and others (2004); 12. Shewry and others (2013); 13. Hidalgo
and others (2009); 14. Suchowilska and others (2009); 15. Suchowilska and others (2012); 16. Zhao and others (2009).
: not available for whole grain flour.
natural genetic resources toward: (1) developing cultivars with sus-
tainable tolerance to both abiotic and biotic stresses, (2) reducing
the risk of loss of biodiversity and foreseeing problems associated
with global climate change, and (3) ensuring the recreation of
ecological integrity as well as sustainable exploitation of biological
diversity.
The most comprehensive comparative investigation to date was
undertaken in Europe (as part of the EU HEALTHGRAIN
project; Shewry and Hey 2015) on 5 wheat species (einkorn, em-
mer, spelt, durum, and bread wheat). The study determined the
total dietary fiber levels in wholeflour of 151 bread wheat cultivars,
10 durum wheat cultivars, and 5 lines of each of einkorn, emmer,
and spelt (Gebruers and others 2008). The agronomic practices
and associated inputs (nutrient and irrigation) have shown rel-
evance to the adaptation of bread wheat plants. A considerable
variation has also been found in the dietary fiber constituents
among the wholegrains of 129 bread wheat genotypes from the
HEALTHGRAIN diversity screen (Andersson and others 2013).
Phytochemicals, vitamins, and antioxidants
In addition to the major components of protein, carbohydrate,
and lipid, wheat grain is an important source of other health-
related ingredients, notably phytochemicals, vitamins, and antiox-
idants as well as macro- and micro-nutrients (Arzani 2011). The
health-promoting vitamins, antioxidants, and phytochemicals of
the ancient wheats have been compared with those of the com-
mon wheat in a number of studies. However, these studies suffer
from being mostly focused on limited numbers of attributes and
genotypes under agronomic conditions favorable to the common
wheat, or from quality assessment impediments as regards samples,
replications, tools, or methods.
Wheat grain is comprised of numerous health-promoting com-
pounds including phenols, which are the most diverse and abun-
dant “phytochemicals” (plant components with biological func-
tions). Polyphenols are comprised of phenolic acids, flavonoids,
and lignans. Recently, they have received considerable attention
because of their potential antioxidant activity and protective ability
to counteract oxidative damage and diseases such as coronary heart
disease, stroke, and cancer (Quinones and others 2013). One of the
major polyphenols present in whole grain wheat products is alkyl-
resorcinol. A comprehensive study by Ziegler and others (2016)
using whole grain flours of 15 genotypes each of einkorn, emmer,
spelt, durum, and bread wheats grown in 4 locations found that
the overall mean concentrations of alkylresorcinol in the species
were comparable, whereas they varied greatly among the geno-
types within each species. On the other hand, the results of the
HEALTHGRAIN study showed higher alkylresorcinol contents
in small-seeded ancient wheats (einkorn, emmer, and spelt) than
in large-seeded modern wheats (durum and bread wheat) (An-
dersson and others 2008; Shewry and Hey 2015). Andersson and
others (2008) reported alkylresorcinol mean values of 581, 595,
and 605 μgg
–1 in the dry matter of emmer, einkorn, and spelt, re-
spectively. Ferulic acid is one of the most important phenolic acids
found in wheat bran which has antioxidant and antiinflammatory
effects. Similar phenolic components including ferulic acid have
been reported for bread wheat and ancient wheats (Abdel-Aal
and Rabalski 2008; Li and others 2008; Serpen and others 2008).
The highest variations among wheat species were reported for
grain carotenoid content. In wheat, the predominant carotenoid
is lutein, which is also the major yellow-colored pigment; only
insignificant amounts of other carotenoids such as β-carotene have
been detected in wheat grain (Abdel-Aal and others 2007). Among
the wheat species, einkorn has been reported to have the highest
lutein content, implying a high antioxidant quality (Abdel-Aal and
others 2007; Abdel-Aal and Rabalski 2008; Lachman and others
2013; Ziegler and others 2016). Einkorn contains up to 10-fold
higher lutein than does bread wheat with durum wheat containing
intermediate amounts of lutein (Ziegler and others 2016). The
higher amounts of carotenoids in durum than in bread wheat
may be ascribed to the selection of yellow-colored grains for the
production of yellow-tinted pasta (Hentschel and others 2002).
Vitamins are classified into water-soluble and fat-soluble com-
pounds. The B vitamin groups comprise various compounds: B1
(thiamine), B2 (riboflavin), B3 (niacin), B6 (pyridoxine), B9 (fo-
late), and B12 (cobalamin) (Gerdes and others 2012). Wheat, its
whole grain in particular, is a valuable source of B1, B2, B3,
B6, and B9 (Shewry and Hey 2015). Folate (B9), also known
as folic acid or folacin, serves numerous functions including its
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Ancient wheats for healthy foods . . .
involvement in the metabolism of protein, formation of red blood
cells, and lowering the risk of neural tube birth defects (Scott
and others 2000). Only folate has been studied in the HEALTH-
GRAIN project using ancient and modern wheat species to ob-
serve minor variations in its concentration, with higher concen-
trations observed in durum (0.74 μgg
–1 dry weight) and emmer
(0.69 μgg
–1 dry weight) than in the other species studied (0.56 to
0.58 μgg
–1 dry weight) (Piironen and others 2008). However, it
should be noted that further studies are required to determine in
detail the amounts and functions of this vitamin to compensate for
the shortcomings of previous studies that focused only on a single
growth environment and on limited numbers of genotypes/species
other than bread wheat.
Wheat grains are also a valuable dietary source of vitamin E,
comprised of 2 tocopherol and tocotrienol groups (each with α-
β-, γ-, and σ- isomers), collectively known as tocochromanols
(Falk and Munne-Bosch 2010). Vitamin E compounds are consid-
ered the most important lipophilic radical-quenching antioxidants
in cell membranes, hence, they are considered vital for human
health (Schneider 2005). Ancient and modern wheat species have
been compared for their tocochromanol contents to detect the
highest total tocol content in einkorn (Lampi and others 2008;
Ziegler and others 2016).
From the above observations, it may be concluded that einkorn
wholegrain has several advantages over the other cultivated an-
cient and modern wheats with higher ploidy levels (tetraploid
and hexaploid). The higher levels of some of the macro- and
micro-nutrients as well as the antioxidant compounds (conju-
gated polyphenols, carotenoids, tocols, alkylresorcinols, and phy-
tosterols) contribute to the excellent nutritional properties of this
diploid ancient wheat.
Other grain components
The high yields of durum and bread wheat cultivars have given
rise to their extensive cultivation and their substantial replacement
for einkorn, emmer, and spelt wheats. However, a growing interest
has more recently been shown in these underutilized crops because
of the advantageous attributes of hulled wheat species such as
their greater amenability to sustainable agriculture and superior
health-related properties over the bread wheat and durum wheat
species (Abdel-Aal and others 1998; Longin and Wurschum 2016).
Hulled wheats can be grown under unfavorable soil and climate
conditions (Arzani 2011). Moreover, they are superior to modern
wheat because of their tolerance to abiotic and biotic stresses such
as diseases, pests, drought, heat, cold, salinity, pollution, and soil
nutrient shortage. In addition they show greater competitiveness
against weeds and tillering ability (Loje and others 2003; Arzani
2011; Hidalgo and Brandolini 2014).
The presence of functional foods in the human diet plays
an important role in both disease prevention and therapy. Al-
though cereal-based products are rich sources of starch, proteins,
and minerals, the presence of antinutritional factors reduces the
bioavailability of these essential nutrients, in particular minerals,
thereby hampering their nutritive values. It is, therefore, crucial
to seek means of counteracting the antinutritional factors in fa-
vor of the nutritional values. Clearly, lowering the antinutrient
(phytate, saponins, tannins, oxalates, and cyanogenic glycosides)
content in the wholemeal wheat flours will lead to enhanced nu-
tritional value of the final products. While the quality of ancient
wheat grain components remains largely unknown, it may be hy-
pothesized that they benefit from less antagonistic interaction(s)
between health-beneficial ingredients and the antinutrients. In
addition, cultivated ancient wheats may contain certain biocom-
pounds with protective functions against insects and pathogens. A
good example is the trypsin/α-amylase inhibitor family found in
the endosperms of ancient wheats that provides defense against an-
imal predators, insects, and bacteria (Fontanini and others 2007).
Such biocompounds can function as immunomodulators or en-
zyme inhibitors.
The modern wheat species (durum and bread wheat) have
been developed from old wheats using modern breeding tech-
niques such as hybridization and subsequent selection of recombi-
nants with high yield potentials. This process has bestowed grain
yield advantages to the modern wheat over its ancestral ancient
wheats. However, not only have some grain quality constituents,
phytochemicals, and nutritional elements declined or been lost
in modern offspring, but some antinutrients have also been en-
hanced. Relatively few long-term, comprehensive studies have
been conducted to determine whether ancient wheat species
grown under traditional (subsistence and low-input) or mod-
ern (high-input) production systems have greater positive health
advantages and lower antinutritional composition than modern
wheat (see review by Shewry and Hey 2015). The evidence from
a decade of work by German researchers (Longin and others 2016;
Longin and W ¨
urschum 2016; Ziegler and others 2015, 2016)
on einkorn, emmer, and spelt wheats as well as those by oth-
ers (D’Antuono and others 1998; Cubadda and Marconi 2002;
Galterio and others 2003; Loje and others 2003; Serpen and oth-
ers 2008; Charmet 2011; Giambanelli and others 2013; Hidalgo
and Brandolini 2014;) have revealed the health benefits of ancient
wheats. Based on a preliminary screening of germplasm in the
field, 15 superior accessions for each of the einkorn, emmer, and
spelt wheats along with 15 commercial cultivars from each of the
durum and bread wheats (to make a total number of 75 geno-
types) were tested at 4 different locations in Germany (Ziegler
and others 2015; 2016; Longin and others 2016). Their results
showed that, although both einkorn and emmer were promising,
einkorn whole grains contained higher minerals and lipophilic
antioxidants (lutein, vitamin E, and steryl ferulates). This suggests
that these properties could be exploited for functional foods and
nutraceutical applications with favorable aroma and color.
Einkorn and emmer wheats are especially promising for the
production of grains with low immunotoxic effects (Gianfrani and
others 2012). However, doubts have been raised as to the safety
of some ancient wheat cultivars for celiac patients (Suligoj and
others 2013), thus suggesting that the immune toxicity for celiac
patients might be genotype-dependent. More recently, serologi-
cal and histological studies have shown that the einkorn cultivar
“Monlis” is toxic to celiac patients, but that it is well tolerated by
the majority of patients with gluten sensitivity (Zanini and others
2015). On the other hand, spelt wheat possessing the D genome
was found to be similar to common wheat in terms of cytotoxicity
as the D genome was found to be related to celiac epitope expres-
sion. Vincentini and others (2007) found no difference between
common and spelt wheats. They, however, noted that the other
ancient wheats were poor in cytotoxic prolamins thereby having
potential as healthy food crops.
The effects of whole grain flour obtained from modern and an-
cient (spelt, emmer, and einkorn) wheats on the development and
course of diabetes in the Zucker diabetic fatty (ZDF) rats were as-
sessed (Thorup and others 2014 ). Results showed that the disorder
was less pronounced with ancient grains fed to the ZDF rats than
with the modern wheat; this was ascribed to a down-regulation
of the expression of important regulatory genes in the liver.
482 ComprehensiveReviews in Food Scienceand Food Safety rVol. 16, 2017 C2017 Institute of Food Technologists®
Ancient wheats for healthy foods . . .
Health-promoting properties of ancient wheat have been cred-
ited to its superior levels of phytochemicals such as carotenoids,
flavonoids, phytosterols, and phenolic compounds (lignans and
ferulic acid) compared with the modern wheat species (Charmet
2011; Giambanelli and others 2013). However, the potential
health-promoting benefits of ancient wheat species await fur-
ther detailed research, particularly under low-input environmental
growing conditions.
Type of Food Consumption and Dietary Diversity
Einkorn, emmer, and spelt possess hulled grains enclosed by
tough glumes (husks) in a semi-brittle rachis. After threshing, the
hulled wheat spikes break into spikelets and uphold the attached
glumes to the grain and the semi-brittle rachis. Either particular
dehullers are used for dehulling or milling/pounding is employed
to remove the glumes from the grains. The ancient wheat is pri-
marily used for human food, but it is also utilized as animal feed.
It is used in making bread and various dishes, some of which have
a long history of cultural dependence, particularly in rural areas
(Hansson 1994; Stallknecht and others 1996). In addition to the
different types of fermented bread, unleavened bread such as cha-
patti and some types of pancake are produced from the ancient
wheat. A porridge is prepared with milk or water or by stirring
in crushed grains, decorticated flour, or extracted starch until it is
fully gelatinized (Hansson 1994; Stallknecht and others 1996).
Einkorn and emmer were initially used as porridge by ancient
civilizations before being used to bake bread. Bread has a long
history dating back to the New Stone Age (Neolithic times) com-
mencing at about 10000 to 8000 BC. Hand-crushed grains mixed
with water were used to bake early bread by laying the dough on
heated stones. The 1st leavened bread was made by people from
Sumeria around 6000 BC by mixing sour dough with unleavened
dough (Belderok 2000). The procedure of preparing bread was
passed down to the Egyptians about 3000 B.C. The Egyptians
started the use of yeast as a potential source of dough rising to
make the bread lighter (Samuel 1996). Nowadays, a wide range
of breads made across the globe come in different sizes, shapes,
textures, colors, crusts, elasticity, eating qualities, and flavors.
Bread made from the emmer wheat flour is widely used in
Switzerland. It is also known as pane di farro in Italy and is available
in some bakeries. Pasta is also produced from emmer wheat in
limited quantities in central Italy at the domestic level but has been
largely rejected by consumers owing to its unappealing texture
(D’Antuono 2013). Bulgur (cracked grains) of emmer is also used
to prepare hard porridge by mixing it with boiled water and
butter. In addition, bulgur whole grains of emmer wheat have
traditionally been used to make different types of soup worldwide.
In some rural areas in Italy and Iran, emmer is used as carbohydrate
source in meals in the same way as rice (Arzani 2011).
Ancient wheat is likely rich in protein, amino acids, antioxidant
compounds, carotenoids, vitamins, and minerals. In combination
with pulses (legumes), it can produce an optimal dietary protein
level for vegetarians, or for those eager to consume a plant-based
food source with a balanced-quality protein.
Ancient and Modern Wheat Flour Blends
Some of the ancient wheats have a distinctive composition such
as resistant starch, carotenoids, phytochemicals, and antioxidants
which offer numerous health benefits. Although ancient wheat
flour may produce a relatively satisfactory loaf of bread, the quality
is not comparable to that made with bread wheat (Arzani 2011;
Brandolini and Hidalgo 2011). Wheat protein is deficient in some
essential amino acids (AA), especially lysine (the 1st limiting AA)
and threonine (the 2nd limiting AA). This naturally results in foods
poor in protein and consequent protein deficiency (Endo and
others 2002; Pichardo and others 2003). Although few studies have
investigated the amino acid composition of ancient wheat proteins,
a wide variation has been observed in the lysine content of einkorn
wheat, some accessions of which have exhibited greater lysine
contents than bread wheat (Loje and others 2003; Hidalgo and
Brandolini 2014). Quantifying the protein content and quality in
ancient and modern wheats, Konvalina and others (2008) observed
a reasonably high amount of protein in the emmer grain; they,
however, found its gluten quality inferior to that of bread wheat.
The poor gluten quality of emmer wheat has been ascribed to
its lack of D genome. Bread-baking with ancient wheat flour
could be more appealing if a blend of flours from ancient and
bread wheats were used. In this way, the high lysine content and
the other health-promoting compounds of ancient wheat could
supplement those of bread flour to achieve a better dietary balance.
Much interest has recently been shown to linking phytonutri-
ents to reducing the incidence of aging-related and chronic hu-
man diseases. It is well established that, from among the numerous
antioxidant compounds found in foods, lipid-soluble antioxidants
play an important role in disease prevention. Curiously, the natural
antioxidant activity of phytonutrients may be associated with their
functional properties contributing to the shelf-life and freshness
of food products. Wheat is an important staple food for a large
proportion of the human population, and it is a source of pro-
tein and energy as well as a source of phytochemical compounds
for human nutrition. As already mentioned, the concentrations of
phytonutrients, however, appear to be lower in bread wheat than
in ancient wheat.
Whole grain flour contains higher amounts of water-insoluble
fibers but lower amounts of the water-soluble ones than white
flour (Ranhotra 1994). Epidemiologists found a strong correla-
tion between low fiber intake, particularly from the gastrointestinal
tract, and many disease states (Birdsall 1985). In developing coun-
tries, there is a general belief among consumers that the greater
plant fiber consumption by people from rural areas protects them
against many diseases such as colon cancer, diverticular disease,
cardiovascular diseases, hemorrhoids, appendicitis, and varicose
veins common to people living in urban areas; in many cases this
has been confirmed experimentally (Anderson and others 2000;
Harris and Kris-Etherton 2010; Liu and others 2015). There is,
however, some ambiguity in inferring a simple link between fiber
intake and cancer risk because food that is rich in fiber may include
other constituents that are linked to cancer prevention. Moreover,
research findings show high-fiber diets to be associated with de-
creased blood pressure both in normal and hypertensive subjects
(Keenan and others 2002; Behall and others 2006). This may be
explained by the fact that a diet enriched with complex carbo-
hydrates enhances glucose metabolism in diabetic individuals by
elevating their insulin sensitivity, leading to reduced insulin dosage
needs (Keenan and others 2002; Lutsey and others 2007; Freeland
and others 2010; Tarini and Wolever 2010). Moreover, a high-
fiber diet is positively associated with the control of obesity and
physical gastrointestinal tract disorders.
High-fiber diets, in particular cereal-based fibers, clearly have
certain health-beneficial effects on diet-related chronic diseases
including obesity, diabetes, cancer, and cardiovascular diseases
(Birdsall 1985; Anderson and others 2000). Accordingly, con-
sumers are able to influence their health by choosing to consume
high-fiber foods including functional cereal products. It should be
C2017 Institute of Food Technologists®Vol.16,2017 rComprehensive Reviews in FoodScience and Food Safety 483
Ancient wheats for healthy foods . . .
noted that cereal product reception by consumers is grounded on
their pleasant texture and flavor which are critical to the devel-
opment of food items. Economic growth and urbanization have
brought about dietary alterations in many countries that favor the
utilization of energy-dense foods high in refined fat and carbo-
hydrates with low dietary fiber (Bach Knudsen and others 2016).
Consumers have replaced whole grain with refined wheat grain
products, with the adverse consequences of reduced dietary fiber,
micronutrient, and phytochemical intakes. As the biochemical
mechanisms of the beneficial health effects of whole grain prod-
ucts are not yet fully understood, it is quite likely that the health
problems stem from the combined action of low dietary fiber and
the inadequacy of a wide variety of phytochemicals (Fardet 2010;
Russell and others 2016). Nevertheless, the lower quantity of fiber
in ancient wheats than in common wheat has been documented
(see Table 1); hence, ancient wheat flour may partly be used as a
substitute for wheat flour in bread and other cereal-based products
to balance the dietary fiber components.
Ancient Wheat for Combating Malnutrition
and Food Insecurity
The growing demand for nutritionally and phytochemically ad-
equate food products has attracted growing attention to ancient
wheat and this interest is likely to continue. One of the ultimate
sustainable goals of wheat improvement is to combat micronutri-
ent malnutrition in developing countries where billions of people
consume staple cereals with inadequate supplies of essential mi-
cronutrients (Bouis 2003; Cakmak 2008). As reported in Table 1
and discussed earlier, ancient wheats, as nutritionally rich species,
are better sources of micronutrients and phytochemicals than are
their modern counterparts (Suchowilska and others 2012; Hidalgo
and Brandolini 2014). To combat the global micronutrient mal-
nutrition, it is, therefore, essential to fortify and enrich staple food
crops with mineral elements as an immediate, but only tempo-
rary solution. International breeding efforts exploit biodiversity
to improve the mineral nutrient levels in food crops. It is well
established that modern high-yielding wheat cultivars have lower
concentrations of mineral nutrients than low-yielding old ones do
(Murphy and others 2008; Shewry and others 2016). Alone, the
germplasm of modern wheat cannot provide the genetic diver-
sity required to develop new wheat cultivars with sufficiently high
grain mineral nutrient concentrations. On the other hand, ancient
wheats, such as einkorn, emmer, and spelt, contain the germplasm
resources that can be exploited to improve wheat grain micronu-
trient value (Genc and McDonald 2008; Suchowilska and others
2012; Hidalgo and Brandolini 2014).
Low-Input/Sustainable Cropping System
Sustainable food production calls for environmentally-friendly
and ecologically sound farming practices. A potentially valuable
measure that may be taken in response to consumer concern for
food quality is to select appropriate crop species such as ancient
wheats through breeding programs and to include them in organic
and sustainable food production systems. Unfortunately, a drastic
climate change is likely to have adverse effects on both yield and
quality of wheat and an effective strategy for the bio-diversification
of wheat varieties and their adaptation to the changing environ-
ment is essential. Global climate change has numerous adverse
effects on food security. Most important are soil degradation and
aggravation of biotic and, in particular, abiotic stresses (Lal 2014;
Arzani and Ashraf 2016). Employing ancient wheat genetic re-
sources as a component of sustainable use of agro-biodiversity
may reduce the risks of climate change to food production. How-
ever, significant enhancements in crop yield stability may equally
be achieved by deploying the less bred/modified species. In the
meantime, such provisions as price adjustments must be made to
compensate for the resulting economic losses to farmers due to
changes in production systems or the use of lower yielding but
nutritionally superior crops. A delicate balance must, therefore, be
struck between the economic outcomes of the new farming prac-
tices at higher costs and a sustainable high-quality crop production.
The focus should be on achieving greater harmony with nature
through enhanced water, soil, and air quality as well as biodiver-
sity. Furthermore, sustainable crop production should encompass
an integrated management of natural resources and the environ-
ment as a whole. Smith and others (2000) noted that intensive
agricultural practices lead to a loss of resilience in the ecosystem
by altering such processes as productivity, nutrient cycling, and
species diversity. The consequences of crop production in this
new environment have an impact not only on farmers but also
millers, bakers, processors, retailers, and consumers. Thus, it is not
solely the farming system that must be in focus.
Public health and environmental concerns in some developed
and developing countries have gradually resulted in a shift from
more intensive cropping practices to more sustainable, low-input
ones. However, one drawback associated with the current high-
yield and high-protein wheat cultivars is that they require high
agrochemical inputs such as nitrogen fertilizers, pesticides, and
herbicides. Ancient wheats that have never been subjected to
breeding programs and their landraces/original forms are presently
available and may provide an important component in the devel-
opment of low input/ high food nutritional food quality systems
to replace the current pattern of high input/ low food nutritional
quality.
The compelling arguments in favor of growing ancient wheat
revolve around reduced agronomic practices, lower energy re-
quirements, environmental sustainability, and higher yield po-
tentials on marginal lands or under organic growing conditions.
They can be produced more sustainably at reduced external in-
puts. Longin and others (2016) evaluated grain yields by early
domesticated and modern wheats (einkorn, emmer, spelt, durum,
and bread wheat) to find that the yields of einkorn, emmer, and
spelt amounted to only 38%, 45%, and 63%, respectively, of the
bread wheat yield. Interestingly, the wheat species were ranked
for their yield potential based on their ploidy levels. In addi-
tion, it was found that the ancient wheats (einkorn, emmer, and
spelt) recorded greater values of plant height (30 cm) than did the
bread and durum wheats. This is consistent with the finding by
Konvalina and others (2010) who evaluated 169 landraces of the
ancient wheat in a field with an annual precipitation of 472 mm
to find that most of the landraces examined showed an inclination
to lodge due to their long and weak stems. This indicates that,
although plant height is genetically determined, the optimum en-
vironmental conditions favorable to the growth of ancient wheat
involve much less external/agronomic inputs, plant density, and
water than do those for the modern wheat.
It is, therefore, conceivable that einkorn with the lowest yield
and ploidy level possesses the highest health benefits (Hidalgo
and Brandolini 2014; Ziegler and others 2016). Being the oldest
cultivated wheat, it was domesticated some 10000 y ago in the
Fertile Crescent. Nevertheless, adaptation of ancient wheats to
the less favorable environmental conditions and lower agronomic
input requirements, combined with their higher tillering ability,
and competitiveness against weeds make them attractive for use
484 ComprehensiveReviews in Food Scienceand Food Safety rVol. 16, 2017 C2017 Institute of Food Technologists®
Ancient wheats for healthy foods . . .
Figure 3–Suggested breeding schemes for (a) introduction of desired traits (health beneficial compounds and tolerance to biotic and abiotic stresses)
from the ancient wheat (einkorn, emmer, and spelt) into modern wheat (durum and bread wheat); and (b) screening of ancient wheat accessions and
subsequent selection of superior accessions (pure line selection) for sustainable grain yield and health benefit traits to develop cultivars for direct
cultivation as sustainable sources of good quality nutrition.
in organic and sustainable production systems. Overpopulation
and acquisition of good-quality lands for industrial development
and urbanization have made it necessary to extend agricultural to
marginal areas. However, the utilizing of marginal lands can only
be worthwhile if adequate yields can be acquired with limited
resources. The cultivation of ancient wheat is particularly impor-
tant in high-altitude marginal lands where they are economically
viable thanks to their cold resistance and low input requirements.
The efficient growth strategies of natural grasses such as ancient
wheat rely on a better partitioning of assimilates between the non-
reproductive and reproductive organs, resulting from a continually
shifting balance between sinks and sources throughout the growing
season. This balance is influenced by abiotic and biotic (pests and
diseases) stresses, particularly under limited water supply.
Diversification of staple crops is recommended not only for in-
creasing the diversity of food sources, but also for establishing a
sustainable, resilient, and food-secure agriculture (Massawe and
others 2016). On the other hand, a narrow focus on only a lim-
ited number of species and cultivars within a species results in an
enormous loss of genetic diversity with associated vulnerability of
ecosystems, extinction of species, and a limited ability of breeders
to respond to future agricultural needs. A major challenge re-
searchers and farmers, in the face of global climate change, is to
save and safeguard genetic biodiversity to allow the necessary adap-
tations to both biotic and abiotic stresses in species of importance.
Conclusions and Future Direction of Research
Although consumers are conscious of the likely diet–health re-
lationships of food, the benefits of diversified cereal grains in the
human diet have not yet been fully recognized. The growing con-
sumption of cereals calls for a profound shift in consumers’ taste
to use breads and products made from wheat types other than
the common wheat. The increasing interest recently shown in
“organic” and “natural” products has led to the “rediscovery” of
ancient wheat on the following grounds:
(i) The capability of both modern and ancient wheats to yield
end-products; this capability could be particularly exploited
to produce many different types of dishes such as whole,
pearled, or broken grains; flour to make bread, pizza, bis-
cuits, and pastries; and semolina to make pasta;
(ii) The nutritional value, high starch-resistance, and healing
properties of the ancient wheat that make it especially useful
for the treatment of such diseases as high blood cholesterol,
colitis, and allergies;
(iii) The ability of the ancient wheat to grow in poor soils with
low input and organic crop systems as well as their higher
tolerance to biotic and abiotic stresses including diseases,
insects, extreme temperature (cold and heat), drought, and
salinity; and
(iv) The putative ancestral and primary gene pool that can be
exploited for the improvement of the health-related traits of
modern wheat (durum and bread wheat).
The potentially dangerous consequences of the genetic ero-
sion of wheat species (Triticum spp. L.) pose a great challenge to
plant domestication and, in particular, to subsequent breeding pro-
grams. Wheat breeders should not only identify the valuable traits
and properties in wheat primitive ancestors in an attempt to intro-
duce them into modern wheat, but should also conduct genetic
improvement projects aimed at their domestication. Two of the 3
ancient wheats, namely emmer and spelt, can readily be used in
hybridization/breeding schemes of durum and bread wheat, while
the emmer wheat genomes can also be combined with goatgrass
(Ae. tauschii) to produce a synthetic hexaploid wheat. The study
conducted by Lage and others (2006) demonstrated that the ge-
netic variation accomplished to improve the quality of tetraploid
emmer wheat could be replicated to produce synthetic hexaploid
wheats.
Ancient wheats are rich in resistant-starch, fiber, minerals, and
phytochemicals. They will, therefore, serve as a valuable source for
improving wheat cultivars with a higher yield and better compo-
sition of health-beneficial compounds. In the primary steps of the
work, assessment of the genetic diversity of the ancient wheats for
their nutritional and health-beneficial properties is recommended,
C2017 Institute of Food Technologists®Vol.16,2017 rComprehensive Reviews in FoodScience and Food Safety 485
Ancient wheats for healthy foods . . .
as has been done by workers on the German Gene Bank collection
for some grain quality traits (see review by Longin and Wurschum
2016). Public–private participations that involve millers and bakers
are suggested to develop ancient wheat products appealing to the
urban consumer’s taste either by blending grain/flour/semolina of
ancient and modern wheats or by utilizing their products inde-
pendently from each other. Diversification in ancient wheat food
products can be improved through process models coming from
European countries like Italy that have successfully produced such
food products.
The prime objective of wheat breeding programs during the
last century was to improve grain yield while also conserving a
certain level of grain protein. However, little attention was paid to
an examination of the nutritional and health-related properties of
grains and their improvement (Leenhardt and others 2006; Arzani
2011). Table 1 compares the whole-grain flour compositions of
einkorn and bread wheats based on the means or ranges of the
genotypes tested within each group. It is necessary to determine
whether both interspecific and intraspecific variations reported re-
late to nutritional properties of the Triticum spp. grains. Figure 3
presents the breeding schemes suggested for the introduction of
traits of merit (health-beneficial compounds and tolerance to bi-
otic and abiotic stresses) from the ancient wheat into the mod-
ern wheat. The proposed schemes also involve screening of an-
cient wheat accessions and their subsequent pure line selection to
achieve sustainable grain yield, enhanced health-promoting com-
pounds, cultivars for propagation and cultivation, and sustainable
sources of good quality nutrition.
The current epidemiological and clinical evidence implies that
a whole grain diet may have a protective function in the initia-
tion and progression of type-2 diabetes (Fung and others 2002;
Montonen and others 2003), coronary heart disease (Jacobs and
others 2002; Behall and others 2006), age-related eye disorders,
and some kinds of cancer (Chatenoud and others 1998; Kasum
and others 2001). The health-enhancing values of whole wheat
grain flour might be attributed to the natural antioxidants, phe-
nolic acids, flavonoids, phytic acids, carotenoids, and tocopherols
(Moore and others 2005; Mpofu and others 2006; Brandolini and
Hidalgo 2011; Lachman and others 2012). Polyphenolic com-
pounds (flavonoids, phenolic acids, and lignans) have a potentially
protective function against oxidative damages such as coronary
heart disease, stroke, and cancer in humans (Quinones and others
2013).
Insufficient data are currently available on the use of ancient
wheat flour as a partial substitute for common wheat flour in
making breads, cookies, and pasta. Although a number of studies
have been carried out on the nutritional and health-related prop-
erties of ancient wheat species, thorough comparisons between
these species and modern wheat will be a complicated task be-
cause of the environment-related aspects. The aspects that need
to be addressed in any such comparative study will have to in-
clude: different agronomic practices and field conditions (such as
plant density and water availability) favorable to each of the groups
and the requirement for reliable phenotypic assessments through
multiple-year and location trials (Shewry and Hey 2015).
The authors declare no conflict of interest.
Acknowledgments
The authors are extremely grateful to P.J.C. Harris, Professor
Emeritus, Univ. of Coventry, U.K., for his critical reading of the
manuscript.
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... About 20% of calories and 55% of carbohydrates are provided by wheat across the globe. Both growth and yield of wheat are negatively influenced by salinity (Arzani and Ashraf, 2017). Wheat bread has high vitamins B, thiamine, and B2riboflavin content (Rakaščan et al., 2019). ...
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2022) Characterization of wheat (Triticum aestivum L.) accessions using morpho-physiological traits under varying levels of salinity stress at seedling stage. Front. Plant Sci. 13:953670. Abiotic stresses are the major stressors affecting wheat (Triticum aestivum L.) production worldwide. The world population is increasing continuously. It is very difficult to feed the population because one-third world's population consumes wheat as a staple food. Among all abiotic stresses, salinity is one that led to a drastic reduction in wheat crop fitness and productivity. Thus, understanding the effects of salinity stress becomes indispensable for wheat improvement programs which have depended mainly on the genetic variations present in the wheat genome through conventional breeding. Therefore, an experiment was conducted using a complete randomized design with four replications, to determine the selection criteria for salinity-tolerant germplasm based on morphophysiological traits at the seedling stage. Three levels of salt solutions, i.e., 4, 8, and 12 dSm −1 were applied and the performance of different genotypes under these three salinities levels was observed. Results depicted that leaf water content and relative water content were correlated with each other. Notably, selection based on these traits increased the performance of other characters. The genotypes G11, G13, G18, G22, and G36 performed best in the salinity stress. So, these genotypes are considered salinity-tolerant genotypes. The genotypes G4, G17, G19, G30, and G38 performed worst in the stress and these were salinity-susceptible genotypes. From the results of the principal component (PC) analysis, the first five PCs were indicated to have a substantial genetic variation from the total of 14 PCs. These PCs showed 75, 73, 65.324, and 65.162% of total variation under normal, salinity level 4, 8, and 12 dSm −1 , Frontiers in Plant Science 01 frontiersin.org Ahmed et al. 10.3389/fpls.2022.953670 respectively. Stomatal conductance, fresh shoot weight and fresh root weight, and dry shoot weight and dry root weight were not significant and negatively associated with all other traits studied, except for relative water and leaf water content. Overall, the results suggested that selection based on leaf water content and relative water content at the seedling stage would genetically improve salinity tolerance. Genotypes with good performance under salt stress conditions may be useful in future breeding programs and will be effective in developing high-yielding salt-tolerant wheat varieties.
... Cluster-und Hauptkomponentenanalysen ergaben schließlich, dass die Anwendung von STI und YSI beim Screening von Wildgenotypen auf Salztoleranz zu einer stabileren Bestimmung der salztolerantesten Genotypen führt. Grains, making up more than 80% of the world's food supply, are threatened by climate change under multiple abiotic stresses, including drought, salinity, and heat (Arzani and Ashraf 2017). Genetic improvement for higher tolerance to abiotic stresses is one of the most economical and sustainable methods to enhance crop productivity and stability (Arzani and Ashraf 2016;Blum 1988;Hussain 2021 6 ). ...
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Salinity is one of the major abiotic factors limiting crop production worldwide. To assess salinity stress tolerance of wild (Hordeum vulgare ssp. spontaneum L.) and cultivated (H. vulgare ssp. vulgare L.) barley genotypes, a two-year field experiment was carried out. Plant height, days to heading, days to anthesis, days to maturity, spike length, grain yield, and yield components were evaluated under normal and salinity stress conditions. The stress stability index (SSI), tolerance index (TOL), yield index (YI), stress tolerance index (STI), geometric mean productivity (GMP), mean productivity (MP), and yield stability index (YSI) were the tolerance indices used in this study. The results showed significant effects of salinity stress and genotype on the measured and calculated variables. The wild genotypes were less affected by salinity stress than the cultivated ones and thus were far superior for the stability indices for yield (STI and YSI). On the other hand, barley cultivars were superior for yield obtained under normal (Yp) and saline (Ys) conditions, MP, GMP, and TOL. A strong relationship was found between grain yield and three indices (STI, YI, and YSI). Ultimately, cluster and principal component analyses suggested that the implementation of STI and YSI in the screening of the wild genotypes for salinity tolerance results in a more robust determination of the most salinity tolerant genotypes.
... Wheat (Triticum aestivum L.) is an important crop, serving as a major source of calories and protein in human diets worldwide (Arzani & Ashraf, 2017). It is consumed in various foods, including pasta, noodles, chapati, naan, bread, and biscuits. ...
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Water stress is one of the major environmental constraints on wheat grain yield worldwide. One way to overcome this limitation is to evolve genetically stress-tolerant wheat genotypes that produce sustainable grain yields in water-scarce conditions. A field experiment was carried out to investigate the genetic diversity of 34 advanced wheat genotypes ( Triticum aestivum L.) and two commercial check varieties (Khirman and TD-1) for grain yield and yield-associated agronomic traits in moisture stress (MS) and well-watered (WW) conditions. Plants were grown in residual moisture in rice fallow land in rainfed conditions without supplementary irrigation, i.e., MS conditions, while two rounds of irrigations were applied for the WW control conditions. Analysis of variance indicated a highly significant ( p < 0.05) variation among genotypes for all the observed agronomic traits in MS and WW conditions. In the MS group, the exotic line IBWSN-1010, mutant line MASR-64, and doubled haploid line DH-12/7 produced the highest grain yield compared to all the contesting wheat genotypes, including check varieties. Grain yield per plot was positively correlated ( r = 0.93) with biological yield per plot in MS conditions. Principal component analysis showed total variations of 21.9%, 20.4%, and 10.1% explained by PC-1, PC-2, and PC-3 in MS, and 22.9%, 14.8%, and 12.1% for PC-1, PC-2, and PC-3 in WW conditions. Our study provides valid information for the selection of newly evolved wheat genotypes and will be useful in future breeding programs.
... In recent years, the scientific interest and appreciation of the quality of traditional wheat varieties has increased for a more sustainable low-input production of grain, as germplasm with an enhanced phytochemical profile, and as a source of adaptive traits in the face of climate change [6,7]. For instance, considering that wheat is a staple food in several countries, anthocyanin-rich grains can be used to produce a wide range of foods with enhanced nutraceutical and pharmaceutical properties. ...
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A main reason of the increasing interest in cereal landraces is their potential to offer more diversified and functional staple food. For instance, landraces are an underexploited resource of pigmented varieties, appreciated for the high accumulation of phytochemicals with known health benefits. This study characterized the chemical, functional, and technological features of the bran, semolina, and grains of two durum wheat (Triticum turgidum L. subsp. durum, Desf.) landraces, named ‘Purple’ and ‘Red’ for their grain color, collected in Ethiopia and grown and sold in southern Italy as a niche product. Specifically, we analyzed the protein content, dry gluten, ash, total polyphenols, anthocyanins, proanthocyanidins, and specific phenolic acids. We also evaluated the antioxidant activity using DPPH- and ABTS-based methods. The two landraces had positive nutritional features, such as a high protein content, a rich and composite range of secondary metabolites (which include specific phenolic acids and anthocyanins), and antioxidant activities in all the fractions analyzed. The germplasm under investigation therefore has a well-justified potential to yield functional products and to diversify durum wheat-based foods.
... It has been well known that the crops most susceptible to mycotoxin contaminations are corn (Park et al. 2018), wheat (Xing et al. 2020, barley (Wan et al. 2020), rice (Ferre 2016), and peanuts (Kang et al. 2015;Yang et al. 2015). According to Food and Agriculture Organization of the United Nations (FAO), wheats are the most produced cereal that are solely aimed for food supply in the world wide (Arzani and Ashraf 2017). The food safety issue regarding wheat and wheat products has become an emerging risk factor that affects the general health status of population, national security, and social economic development for any country or nation. ...
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Abnormal climate changes have resulted in over-precipitation in many regions. The occurrence and contamination levels of mycotoxins in crops and cereals have been elevated largely. From 2017 to 2019, we did investigation targeting 15 mycotoxins shown in the wheat samples collected from Shandong, a region suffering over-precipitation in China. We found that deoxynivalenol (DON) was the dominant mycotoxin contaminating wheats, with detection rates 304/340 in 2017 (89.41%), 303/330 in 2018 (91.82%), and 303/340 in 2019 (89.12%). The ranges of DON levels were < 4 to 580 μg/kg in 2017, < 4 to 3070 μg/kg in 2018, and < 4 to 1540 μg/kg in 2019. The exposure levels were highly correlated with local precipitation. Male exposure levels were generally higher than female’s, with significant difference found in 2017 (1.89-fold, p = 0.023). Rural exposure levels were higher than that of cities but not statistically significant (1.41-fold, p = 0.13). Estimated daily intake (EDI) and margin of exposure (MoE) approaches revealed that 8 prefecture cities have probabilistically extra adverse health effects (vomiting or diarrhea) cases > 100 patients in 100,000 residents attributable to DON exposure. As a prominent wheat-growing area, Dezhou city reached ~ 300/100,000 extra cases while being considered as a major regional contributor to DON contamination. Our study suggests that more effort should be given to the prevention and control of DON contamination in major wheat-growing areas, particularly during heavy precipitation year. The mechanistic association between DON and chronic intestinal disorder/diseases should be further investigated. Graphical abstract
... The range of temperatures has determined diversity of cultivated vegetables and fruits in Iraq. Wheat cultivation dominates in the North of Iraq given the origin of wheat from the Fertile Crescent region in northern Iraq [15]. In the South of Iraq, date fruits are mainly cultivated and believed to have originated from this region [16]. ...
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The tradition of making fermented foods and beverages in Iraq dates back to 7500 BC. These fermented foods and beverages are represented by meat-, milk-, vegetable-, and fruit-based products reflecting diversity of agricultural production in ancient Iraq (Mesopotamia). Although the recipes for some fermented foods and beverages were lost throughout history, those remaining foods and beverages occupy a noticeable position in modern Iraqi cuisine. In this review, knowledge and techniques for preparation of 5 traditional fermented foods, i.e. Basturma, Smoked Liban, Aushari cheese, Turshi, and Sour Khobz, and 3 fermented beverages, i.e. Shanina, Sharbet Zbeeb, and Erk Sous in Iraq, are documented. Traditional fermented foods and beverages have multiple health benefits because of high content of probiotics and bioactive compounds. Traditional fermented foods and beverages are made using the back-slopping technique which ensures safety of production and maintains organoleptic properties. The review highlights the potential of fermented foods and beverages for their large-scale commercialization.
... Einkorn is cultivated recently in some poor soils of Southern Europe, Minor Asia, Caucasus, North Africa (Miroshnichenko et al. 2017). As a functional food, this diploid cereal has many benefits over the cultivated modern tetraploid and hexaploid wheat varieties, such as higher level of macro-and micronutrients (phosphorus, sulfur, magnesium, zinc, manganese etc.) and various antioxidant (conjugated polyphenols, carotenoids, tocols, alkylresorcinols, and phytosterols) compounds (Suchowilska et al. 2012;Zaharieva and Monneveux 2014;Arzany and Ashraf 2017). Furthermore, einkorn can be used in modern wheat breeding programmes as a source of new traits for pest and disease resistance and abiotic stress tolerance (Zaharieva and Monneveux 2014;Longin et al. 2016;Miroshnichenko et al. 2017). ...
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Einkorn ( Triticum monococcum L.) can be applied as a model species for cereal genomic studies due to its small genome size and high level of polymorphism. The in vitro somatic tissue culture protocol in einkorn was significantly improved recently, however the in vitro androgenesis remained an unresolved research topic. Five different pre-treatments were compared to study the effects of stress pre-treatments on the efficiency of androgenesis in two einkorn genotypes. The long cold pre-treatment (2 weeks, 4 °C) of donor tillers increased significantly the number of microspore derived embryo-like structures (ELS). Green and albino plantlets were regenerated from these structures. The ploidy level of microspore-derived green plantlet was determined as haploid by flow cytometric analyses. This is the first report published on the successful androgenesis induction (ELS production) and green- and albino plantlet regeneration in in vitro anther culture of the recalcitrant einkorn wheat ( Triticum monococcum L.).
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Wheat grain quality characteristics have experienced increasing attention as a central factor affecting wheat end-use products quality and human health. Nonetheless, in the last decades a reduction in grain quality has been observed. Therefore, it is central to develop efficient quality-related phenotyping tools. In this sense, one of the most relevant wheat features related to grain quality traits is grain nitrogen content, which is directly linked to grain protein content and monitorable with remote sensing approaches. Moreover, the relation between nitrogen fertilization and grain nitrogen content (protein) plays a central role in the sustainability of agriculture. Both aiming to develop efficient phenotyping tools using remote sensing instruments and to advance towards a field-level efficient and sustainable monitoring of grain nitrogen status, this paper studies the efficacy of various sensors, multispectral and visible red-green-blue (RGB), at different scales, ground and unmanned aerial vehicle (UAV), and phenological stages (anthesis and grain filling) to estimate grain nitrogen content. Linear models were calculated using vegetation indices at each sensing level, sensor type and phenological stage. Furthermore, this study explores the up-scalability of the best performing model to satellite level Sentinel-2 equivalent data. We found that models built at the phenological stage of anthesis with UAV-level multispectral cameras using red-edge bands outperformed grain nitrogen content estimation (R² = 0.42, RMSE = 0.18%) in comparison with those models built with RGB imagery at ground and aerial level, as well as with those built with widely used ground-level multispectral sensors. We also demonstrated the possibility to use UAV-built multispectral linear models at the satellite scale to determine grain nitrogen content effectively (R² = 0.40, RMSE = 0.29%) at actual wheat fields.
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Background Septoria tritici blotch (STB), caused by Zymoseptoria tritici ( Z. tritici ) , is an important biotic threat to durum wheat in the entire Mediterranean Basin. Although most durum wheat cultivars are susceptible to Z. tritici , research in STB resistance in durum wheat has been limited. Results In our study, we have identified resistance to a wide array of Z. tritici isolates in the Tunisian durum wheat landrace accession ‘Agili39’. Subsequently, a recombinant inbred population was developed and tested under greenhouse conditions at the seedling stage with eight Z. tritici isolates and for five years under field conditions with three Z. tritici isolates. Mapping of quantitative trait loci (QTL) resulted in the identification of two major QTL on chromosome 2B designated as Qstb2B_1 and Qstb2B_2 . The Qstb2B_1 QTL was mapped at the seedling and the adult plant stage (highest LOD 33.9, explained variance 61.6%), conferring an effective resistance against five Z. tritici isolates. The Qstb2B_2 conferred adult plant resistance (highest LOD 32.9, explained variance 42%) and has been effective at the field trials against two Z. tritici isolates. The physical positions of the flanking markers linked to Qstb2B_1 and Qstb2B_2 indicate that these two QTL are 5 Mb apart. In addition, we identified two minor QTL on chromosomes 1A ( Qstb1A ) and chromosome 7A ( Qstb7A ) (highest LODs 4.6 and 4.0, and explained variances of 16% and 9%, respectively) that were specific to three and one Z. tritici isolates, respectively. All identified QTL were derived from the landrace accession Agili39 that represents a valuable source for STB resistance in durum wheat. Conclusion This study demonstrates that Z. tritici resistance in the ‘Agili39’ landrace accession is controlled by two minor and two major QTL acting in an additive mode. We also provide evidence that the broad efficacy of the resistance to STB in ‘Agili 39’ is due to a natural pyramiding of these QTL. A sustainable use of this Z. tritici resistance source and a positive selection of the linked markers to the identified QTL will greatly support effective breeding for Z. tritici resistance in durum wheat.
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Human history was transformed with the advent of agriculture in the Fertile Crescent with wheat as one of the founding crops. Although the Fertile Crescent is renowned as the center of wheat domestication, archaeological studies have shown the crucial involvement of Çatalhöyük in this process. This site first gained attention during the 1961-65 excavations due to the recovery of primitive hexaploid wheat. However, despite the seeds being well preserved, a detailed archaeobotanical description of the samples is missing. In this article, we report on the DNA isolation, amplification and sequencing of ancient DNA of charred wheat grains from Çatalhöyük and other Turkish archaeological sites and the comparison of these wheat grains with contemporary wheat species including T. monococcum, T. dicoccum, T. dicoccoides, T. durum and T. aestivum at HMW glutenin protein loci. These ancient samples represent the oldest wheat sample sequenced to date and the first ancient wheat sample from the Middle East. Remarkably, the sequence analysis of the short DNA fragments preserved in seeds that are approximately 8400 years old showed that the Çatalhöyük wheat stock contained hexaploid wheat, which is similar to contemporary hexaploid wheat species including both naked (T. aestivum) and hulled (T. spelta) wheat. This suggests an early transitory state of hexaploid wheat agriculture from the Fertile Crescent towards Europe spanning present-day Turkey.
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Fibroblast growth factor 21 (FGF21) is the first known endocrine signal activated by protein restriction. Although FGF21 is robustly elevated in low-protein environments, increased FGF21 is also seen in various other contexts such as fasting, overfeeding, ketogenic diets, and high-carbohydrate diets, leaving its nutritional context and physiological role unresolved and controversial. Here, we use the Geometric Framework, a nutritional modeling platform, to help reconcile these apparently conflicting findings in mice confined to one of 25 diets that varied in protein, carbohydrate, and fat content. We show that FGF21 was elevated under low protein intakes and maximally when low protein was coupled with high carbohydrate intakes. Our results explain how elevation of FGF21 occurs both under starvation and hyperphagia, and show that the metabolic outcomes associated with elevated FGF21 depend on the nutritional context, differing according to whether the animal is in a state of under- or overfeeding.
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Einkorn, emmer, and spelt are old wheat species that have fed the world for centuries before they have nearly completely been replaced by modern bread wheat. Nowadays, the diversity of these old species lies frozen in gene banks and rare attempts aim to exploit them as a source for genetic diversity in modern wheat breeding. Here, we want to raise a debate on a more holistic exploitation of ancient species via their direct introduction to the consumer market as high quality products. Although exemplified only for ancient wheat species, this innovative self-financing strategy can be directly extended to other species. A central requirement for this concept is intensive communication, coordination, and interdisciplinary research along the entire production chain from farm to fork.
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The prediction is that food supply must double by 2050 to cope with the impact of climate change and population pressure on global food systems. The diversification of staple crops and the systems in which they grow is essential to make future agriculture sustainable, resilient, and suitable for local environments and soils. http://authors.elsevier.com/a/1Sxzd4rGdj5szP
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Bread and flour-based foods are an important part of the diet for millions of people worldwide. Their complex nature provides energy, protein, minerals and many other macro- and micronutrients. However, consideration must be taken of three major aspects related to flour and bread. The first is that not all cultures consume bread made from wheat flour. There are literally dozens of flour types, each with their distinctive heritage, cultural roles and nutritive contents. Secondly, not all flours are used to make leavened bread in the traditional (i.e., Western) loaf form. There are many different ways that flours are used in the production of staple foods. Thirdly, flour and breads provide a suitable means for fortification: either to add components that are removed in the milling and purification process or to add components that will increase palatability or promote health and reduce disease per se. Flour and Breads their Fortification in Health and Disease Prevention provides a single-volume reference to the healthful benefits of a variety of flours and flour products, and guides the reader in identifying options and opportunities for improving health through flour and fortified flour products. Examines those four and break related agents that affect metabolism and other health-related conditions. Explores the impact of compositional differences between flours, including differences based on country of origin and processing technique. Includes methods for analysis of flours and bread-related compounds in other foods.