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Sesame: Bioactive Compounds and Health Benefits

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DOI: 10.1007/978-3-319-78030-6_59

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Sesame: Bioactive Compounds and Health
Benefits

Niti Pathak, Asani Bhaduri, and Ashwani K Rai

Contents
1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Bioactive Compounds in Sesame . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2.1 Phenolics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
2.2 Minor Phenolics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
2.3 Phytosterols . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
2.4 Phytates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
2.5 Polyunsaturated Fatty Acids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
2.6 Short-Chain Peptides, Protein Hydrolysates, and Their Functional Properties . . . . . . . 11
3 Quantitative Insight in Lignan and Tocopherol Content: Interspecific and Intraspecific
Variation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
4 Biotechnological Approaches for Sesame . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
4.1 Genome Sequencing of Sesame Crop: Unraveling the Oil Biosynthetic Pathway . . . 12
4.2 Functional Gene Expression: “Lignan Biosynthetic Genes” . . . . . . . . . . . . . . . . . . . . . . . . . . 13
5 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

Abstract
Sesame is a valuable oilseed crop that contains various nutritionally rich bioactive
compounds including lignans, tocopherol homologues, phytosterols, etc. Lignans
are the product of oxidative coupling of β-hydroxyphenylpropane. Sesame has a

N. Pathak
Department of Botany, Deshbandhu College, University of Delhi, Delhi, India
e-mail: [email protected]
A. Bhaduri
Cluster Innovation Centre, University of Delhi, Delhi, India
e-mail: [email protected]
A.K. Rai (*)
Department of Botany, Banaras Hindu University, Varanasi, India
e-mail: [email protected]

# Springer International Publishing AG 2018 1

J.-M. Mérillon, K.G. Ramawat (eds.), Bioactive Molecules in Food, Reference Series in
Phytochemistry, https://doi.org/10.1007/978-3-319-54528-8_59-1
2 N. Pathak et al.

combination of glycosylated lignans and oil-dispersed lignans. Based on their

medicinal and pharmacological properties, the most important lignans are
sesamin, sesamol, sesamolin, and sesaminol. Tocopherols (vitamin E compounds)
are the lipid-soluble free radicals and constitute a major part of human diet. In
sesame seeds, α-, γ-, and δ-tocopherols are found as tocopherol homologues. In
addition to lignans and tocopherols, sesame is an important source of phytosterols,
phytates, polyunsaturated fatty acids, and bioactive peptides. However, utilization
potential of many of these compounds has not yet been fully understood. This
chapter delves into the presence of multifarious bioactive components in sesame
seeds, their biosynthetic pathway, and functional importance.

Keywords
Bioactive compounds
Sesame
Sesamin
Sesamolin
Sesamol
Pinoresinol

Tocopherol
Phytosterols

Abbreviations
CYP81Q1 Sesamin synthase
DIR1 Dirigent protein
DMPQ 2,3-Dimethyl-5-phytyl-1,4-hydroquinol
VTE1 Tocopherol cyclase
γ-TMT γ-Tocopherol methyltransferase

1 Introduction

Sesame (Sesamum indicum, family Pedaliaceae) is considered as one of the earliest

domesticated crops and oilseed plants known to mankind with its multifarious uses.
It is found in the tropics and subtropics but is most common in the narrower belt
closer to the equator, mostly north of it [1]. Basically, sesame is a crop of the
developing countries in more southern latitudes. For a long time, it is being used
in religious rituals in India, Egypt, and Persian region [2, 3]. Sesame is regarded as
“queen of oilseeds” because of its oil quality [4, 5], sterols, and antioxidative agents,
i.e., methylenedioxyphenyl compounds, sesamin, sesamolin, and tocopherols that
act as nutraceuticals and impart resistance to oil against oxidative deterioration.
Further, the composition of dry decorticated sesame seed per 100 g includes edible
portion, i.e., water (3.75 g), energy (2640 kJ; 631 kcal), protein (20.45 g), fat
(61.21 g), carbohydrate (11.73 g), dietary fiber (11.6 g), high amounts of Ca
(60 mg), Mg (345 mg), P (667 mg), K (370 mg), Fe (6.36 mg), Zn (6.73 mg),
vitamin A (66 IU), thiamin (0.70 mg), riboflavin (0.09 mg), niacin (5.80 mg), folate
(115 μg), alpha-tocopherol (1.68 mg), and no ascorbic acid [6]. The presence of
oxalic acid in sesame seeds makes it little bitter in taste.
The sesame oil is comprised of 83–90% unsaturated fatty acids that contain
glycerides of oleic acid (36–54%) and linoleic acid (38–49%). Other components
Sesame: Bioactive Compounds and Health Benefits 3

Fig. 1 Seed color variation in two sesame species cultivated in India

are saturated fatty acids (myristic acid, 0.1% or less; palmitic acid, 8–12%; stearic
acid, 3.5–7%; arachidonic acid, 0.5–1%). The unsaponifiable matter (1.2%) includes
tocopherols and the lignans sesamin (0.1–0.6%), sesamolin (0.25–0.3%), sesamol,
and sesaminol, which give the oil its resistance to rancidity. Extracted sesame cake
varies in color from light yellow to grayish black, depending on the dominant seed
coat color. Its chemical composition also varies according to cultivar, method of oil
extraction, and the presence of testa. The sesame cake has ample amount of calcium
and phosphate; protein content ranges from 35% to 47% but is deficient in lysine.
Sesame seed color show variation in different species which could partly be
attributed to changes in composition of bioactive phenolics i.e., lignans and tocoph-
erols (Fig. 1). The present chapter provides detailed and updated information on the
bioactive components present in sesame seeds, their importance, and health benefits.
In particular, sesame lignans and tocopherols have been focused at length: the
biosynthetic pathway, the gene regulation, and the biotechnological approaches to
enhance the concentration of lignans and tocopherols in sesame crop.

2 Bioactive Compounds in Sesame

Bioactive compounds are beneficial components in food and are accountable for
disease-preventing properties. They include a range of chemical compounds includ-
ing phenolics, carotenoids, phytosterols, and polyunsaturated fatty acids. These
compounds are often utilized as antioxidants and other purposes such as inhibiting
cholesterol absorption, blocking the activity of bacterial toxins, etc. Our recent
review on value addition in sesame crop has elaborated the production of value-
added products such as sesame oil and meal, thereby enhancing its economic
importance [7]. The target candidates for this study are bioactive compounds
such as lignans, tocopherols, phytates, phytosterols, and polyunsaturated fatty
acids (Fig. 2).
4 N. Pathak et al.

Fig. 2 Distribution of bioactive components in sesame seeds

2.1 Phenolics

Natural phenolic compounds are secondary metabolites, which are widely distrib-
uted in the plant kingdom. Plant phenols are noted for their role in prevention of
various ailments associated with oxidative stress such as cardiovascular and neuro-
degenerative diseases and cancer [8]. Phenolics are characterized by the presence of
aromatic ring (at least one) coupled with a few hydroxyl groups. The antioxidative
potential of phenolics is very high as these compounds can produce stable radical
intermediates utilizing electrons. Due to their antioxidative potential, they play an
important role in the stabilization of edible oils and protection from off-flavor
formation [8]. Recent findings suggest that phenolics could play an important role
in conditions associated with oxidative stress [9, 10]. The antioxidant property of
sesame seed and its oil along with various health properties is attributed to the
presence of lignans such as sesamin, sesamolin, sesaminol, sesangolin, 2-episalatin,
and tocopherol isomers [11]. Sesame oil is extremely resistant to oxidative rancidity
due to the presence of chemically related compounds sesamol and sesamol dimer.

2.1.1 Lignans: Chemistry and Biosynthesis

The common feature of many natural products is recognized as a C6C3 unit, i.e., a
propylbenzene or phenylpropanoid skeleton [12]. In a review of natural resins,
Sesame: Bioactive Compounds and Health Benefits 5

Haworth was first to suggest (1936) [13] that the class of compounds derived from
two C6C3 units having β,β0 linkage (8–80 bond) should be called as lignans. Lignan is
actually a constituent of lignin, a generic name for the compound resulting from two
p-hydroxyphenylpropane molecules. Sesame seed contains two major groups of
lignans: (1) oil-soluble lignans (sesamin, sesamolin, sesaminol, sesamolinol, and
pinoresinol) and (2) glycosylated water-soluble lignans (sesaminol triglucoside,
pinoresinol triglucoside, sesaminol monoglucoside, pinoresinol monoglucoside,
and two isomers of pinoresinol diglucoside and sesaminol diglucoside) [14–16].
The antioxidative property of sesame seed is associated with lignan components
present in the oil and are unique to sesame. The lignans, sesamin, and sesamolin and
their derivatives prevent oxidation of the oil and give it a long shelf life and stability
[17]. Lignans and tocopherols are also reported to act in coordination resulting in
enhanced vitamin E activity [18]. Namiki [19] suggested that the bioactive com-
pounds present in sesame seed cannot individually describe the high oxidative
stability of roasted sesame oil, but the cumulative effect of all the components of
sesame oil protects the roasted sesame oil from oxidative deterioration.
The oil fraction of most oilseeds comprises mainly of triacylglycerols (95–99%)
acting as a dispersing media for a wide range of lipophilic and amphiphilic second-
ary metabolites, together known as the unsaponifiable fraction of the seed. These
unsaponifiable portions can be extracted with lipophilic solvents after saponification
of acyl lipids [20] and are of importance as the presence of the lignans may be taken
as marker for genuineness of the oil. Other constituents, tocopherols and
tocotrienols, are strong antioxidants and provide stability to the oil. Also, lignans,
especially sesamin, have currently been recognized for possessing interesting phys-
iological bioactivities against chronic diseases.
Sesaminol, a water-soluble glycoside along with fat-soluble sesamin and
sesamolin, constitutes the major lignans of sesame seeds. The isomer of sesamin
called episesamin is generated in the process of refining sesame oil. The functional
methylenedioxyphenyl group of most of these lignans especially sesamin,
sesamolin, and sesangolin initiates the activities [21, 22], and these molecules in
turn utilize their effect via inhibition of liver microsome oxidases [23].

Sesamin Biosynthesis
Phenylalanine and tyrosine are precursors for various plant substances such as
tannins, polymeric lignin, and lignans. Phenylalanine is converted to cinnamic
acid by the enzyme phenylalanine ammonia lyase (PAL). This is followed by a
series of hydroxylation, methylation, and reduction leading to production of
coumaric acid, caffeic acid, ferulic acid, and eventually E-coniferyl alcohol [24].
However, it has also been claimed that [14C]-labeled tyrosine is incorporated into
sesamin when administered to the cell suspension cultures of Sesamum indicum [25].
E-coniferyl alcohol has also been shown to undergo stereoselective coupling
to synthesize (þ)-pinoresinol in Sesamum indicum seeds (Fig. 3). (þ)-Pinoresinol
is then metabolized further, and methylenedioxyphenyl group is added in matur-
ing sesame seeds which result in production of (þ)-piperitol and (þ)-sesamin and
(þ)-sesamolin [26]. A 78 kDa dirigent protein is known to assist in a stereoselective
6 N. Pathak et al.

Fig. 3 Diagrammatic representation of sesamin biosynthetic pathway depicting conversion of

coniferyl alcohol to pinoresinol with unstable intermediate piperitol followed by conversion to
sesamin. Sesamin is further converted to sesamolin. Boxes indicate action of enzymes

bimolecular coupling to produce (þ)-pinoresinol [27]. Lignan synthesis is develop-

mentally regulated and depends upon the stage of seed maturity. The most mature
seeds of 8 weeks efficiently convert (þ)-pinoresinol into (þ)-piperitol and (þ)-
sesamin, while younger seeds have higher conversion to (þ)-sesamolin [28].
Recently, it is shown that the synthesis of sesamin from pinoresinol is catalyzed
by CYP81Q1 in two steps [29].

2.1.2 Tocopherol Chemistry and Biosynthesis

Tocochromanols have both the hydrophobic and hydrophilic elements – these bio-
molecules generally have a lipophilic isoprenoid side chain linked to the membrane
lipids and a polar chromanol ring toward the membrane surface. Tocochromanols
inhibit membrane lipid peroxidation and scavenge reactive oxygen species. It is a
well-known fact that antioxidants neutralize free radicals, thereby preventing DNA
damage. Being scavenger of reactive oxygen species, tocopherols minimize free
radical attack and interrupt lipid peroxidation. In this way, these molecules protect
cell membranes, enable lipid repair and replacement, and are useful in preventing
cancer and heart diseases [30]. Tocopherols are known to play a role in plant
metabolism, for instance, sugar transport from leaves to phloem [31].
Tocopherols are an important group of plant phenolics that have antioxidative
activity and nutritional values [32]. They belong to a family of molecules that have a
chromanol ring (chroman ring with an alcoholic hydroxyl group) and a 12-carbon
Sesame: Bioactive Compounds and Health Benefits 7

aliphatic side chain containing two methyl groups in the middle and two more
methyl groups at the end. Plants synthesize eight different vitamin E forms, includ-
ing α-, β-, γ-, and δ-tocopherols and α-, β-, γ-, and δ-tocotrienols [33]. All tocoph-
erols and tocotrienols consist of a chromanol ring and varying number of methyl
groups on the chromanol ring. Metabolic fate and biological activities of tocols
depend upon their structural features. The entire isoforms act as lipid antioxidants,
and α-tocopherol has the highest vitamin E activity [34, 35]. The difference between
tocopherols and tocotrienols is that tocopherols have a saturated tail, while
tocotrienols have an unsaturated tail.
Higher plants like dicots have tocopherols in almost all parts including roots,
stems, leaves, flowers, fruits, and seeds [36, 37]. However, the total tocopherol
content and different forms of tocopherols in these tissues differ considerably.
Among the tocopherols, α-tocopherol is the predominant form in photosynthetic
tissues such as stems and leaves. In most seed crops, α-tocopherol is present only in a
minor form, and γ- and δ-tocopherols tend to predominate [38].

Tocopherol Biosynthesis
The hydroquinone ring of tocopherol is derived from the shikimate pathway
of aromatic amino acid synthesis (Fig. 4). Biosynthesis of homogentisate, the
precursor for tocopherol, tocotrienol, and plastoquinone, is catalyzed by p-
hydroxyphenylpyruvate dioxygenase (HPPD) [39]. After attachment of the hydro-
phobic side chain by homogentisate phytyl transferase (HPT1/VTE2) [40, 41] and on
methylation (VTE3) [42, 43], 2,3-dimethyl-5-phytyl-1,4-hydroquinol (DMPQ) is
formed, which is converted to γ-tocopherol by tocopherol cyclase (VTE1) [44, 45].
Tocopherol cyclase is considered as a key enzyme in tocopherol biosynthesis. Final
methylation by γ-tocopherol methyltransferase (γ-TMT, VTE4) results in the pro-
duction of α-tocopherol.

2.1.3 Sesame Seed: Lignan and Tocopherol Bioactivities

Sesamin and sesamolin were initially considered as the major lignans in sesame
seeds [46]. Sesaminol, another major lignan, was identified later from sesame seeds
[47]. These seed components possess unique properties, and regular human con-
sumption helps in lowering blood lipids [48] and arachidonic acid levels [49]. The
sesame seed lignans are also known to reduce cholesterol level by inhibiting its
absorption and synthesis simultaneously [50]. These lignans are anticarcinogenic
[51] and anti-inflammatory [52] and are known to increase hepatic fatty acid
oxidation [53]. They have immunomodulatory activities [54] and possess antihyper-
tensive [55, 56] and neuroprotective effects against hypoxia or brain damage [57].
In addition to lignans, multiple tocopherol homologues [α-tocopherol (αT), δ-
tocopherol (δT), and γ-tocopherol (γT), tocotrienols] are also present in sesame
seeds, which possess antioxidative and health-promoting properties. Sesame lignans
are found to exhibit synergistic effect with tocopherols on vitamin E activity and act
as specific inhibitor of fatty acid metabolism in humans [58]. These nutritional
properties of the sesame seeds contributed by the principal factors lignans and
tocopherols have promoted their use in the daily diet worldwide.
8 N. Pathak et al.

Fig. 4 Diagrammatic representation of tocopherol biosynthetic pathway depicting conversion of p-

hydroxyphenylpyruvate to homogentisate, which gets converted to methylated hydroquinol and
finally to tocopherol homologues

Sesame lignans have antioxidant and tocopherol-sparing activities [59–62]. They are
reported to reduce cholesterol level [48, 50, 63] and exhibit antihypertensive [64] and
anti-inflammatory activities [65] as well as affect lipid metabolism by enhancing gene
expression and hepatic enzyme (acyl CoA oxidase, carnitine palmitoyltransferase,
bifunctional enzyme, and 3-ketoacyl-CoA-thiolase) activities involved in fatty acid
oxidation [66, 67]. On the other hand, lignans reduce the activities of enzymes involved
in lipogenesis (acetyl-CoA carboxylase, fatty acid synthase, ATP citrate lyase, glucose-
6-phosphate dehydrogenase, and pyruvate kinase) by altering the gene expression [66].
Hence, sesame plays a crucial role in minimizing the vulnerability and increasing the
safety against atherosclerosis, cancer, and cardiovascular diseases [48, 64, 68].
Thus, sesame has both preventive and therapeutic values in a variety of chronic
diseases, owing to its rich lignan content and its antioxidative, anticholesterolemic,
and antihypertensive properties. Sesame lignans, especially sesamin, are absorbed in
human body, undergo enterohepatic circulation, and are converted into strong
antioxidative metabolites. Intestinal bacteria metabolize them into other bioactive
compounds such as mammalian lignans enterolactone and enterodiol compounds,
which the role in breast, prostate, and colon cancers, bone health, and cardiovascular
diseases is understudy [69–71].
In traditional Chinese and Indian systems of medicine like Ayurveda, sesame
occupies an important position with diverse pharmaceutical application. In China,
Sesame: Bioactive Compounds and Health Benefits 9

sesame oil is used in treatment of toothaches and gum diseases [72]. In India, it is
being used as an antibacterial mouthwash to relieve anxiety and insomnia and in the
treatment of blurred vision, dizziness, and headache [72]. Sesame oil is also noted
for burn-healing effect [73]. Moist-exposed burn ointment (MEBO) a purely herbal
formulation from China contains sesame oil as an active ingredient along with β-
sitosterol, berberine, and other plant extracts in trace amount [74] and is usually used
in managing the burns of the face, neck, and hand [73].
The main antioxidant activity of α-tocopherol is to break the radical chain in
membranes and lipoproteins [75]. Its antioxidant potential along with various func-
tions at the molecular level mitigates the possibility of cardiovascular diseases and
cancers [76, 77]. Other tocols that are present in lesser amount can also perform the
antioxidative and biological activities. Gamma-tocopherol (γ-T), for instance, is
more potent in decreasing platelet aggregation and LDL oxidation and delaying
intra-arterial thrombus formation than that of α-tocopherol [78, 79]. Tocotrienols
inhibit cholesterol biosynthesis [80] and are found effective in reducing the tendency
of breast cancer [81]. Thus, tocopherols have high antioxidant, antitumor, and
hypocholesterolemic potential. γ-Tocopherol is the major tocopherol in sesame,
whereas α- and δ-tocopherols are present in very small amounts. It has been reported
that γ-tocopherol is a more effective antioxidant compared to other tocopherols [82]
but has poor vitamin E activity in biological systems [76].
Tocopherols terminate the recyclable chain reaction of polyunsaturated fatty acid
(PUFA) radicals produced by oxidation of lipid [83]. The lipid peroxy radicals that
are scavenged by tocopherols are converted into tocopheroxyl radicals, which with
the help of ascorbate and other antioxidants are recycled back as the corresponding
tocopherol [84]. In this manner, a tocopherol molecule can undergo in multiple lipid
peroxidation chain-breaking events before it is finally degraded.

2.2 Minor Phenolics

In addition to tocopherols and lignans, sesame contains other phenolics like

naphthoquinone and phenolic acids in trace amounts [85–88]. Sesamol, which is
known as a strong free radical scavenger [89], is also present in sesame oil. It is a free
phenol with methylenediphenoxy group.

2.3 Phytosterols

Phytosterols (sterols and stanols) are plant triterpenes with preventive functions
in many diseases especially cancer. They have been shown to possess antioxidant
[90], anti-inflammatory [91], and antibacterial properties [92]. Similar in structure
to cholesterol (phytosterols have extra methyl group at C-24 position), these plant
sterols, when digested, compete with cholesterol for small intestine absorption leading
to lowering of the cholesterol level in blood [93]. The recommended functional foods
usually contain phytosterols extracted from plant sources, or at times processed foods
10 N. Pathak et al.

have phytosterols as supplement and are sold as cholesterol-lowering foods. Although

corn and legumes are used to extract phytosterols, it is the sesame seeds that have the
highest (400–413 mg 100 g1) amount of phytosterols [94].
In a recent study, Gharby et al. [95] noted that β-sitosterol constitutes a major
portion of phytosterols in sesame seed and oil, and campesterol and stigmasterol are
the other important sterols present. Compared to other phytosterols, β-sitosterol has
been studied more extensively for its beneficial and physiological effect on human
being. β-Sitosterol lowers cholesterol level [96], enhances immunity, and has anti-
inflammatory properties [97]. The other major component is campesterol, which
accounts for about 17.8% of the total sterols. Stigmasterol and Δ5-avenasterol
measure about 6.4% and 10.2%, respectively, in sesame oil. Minor sterols present
are Δ7-stigmasterol and Δ7-avenasterol. The total sterol content in sesame seed oil is
approximately 540 mg/100 g oil.

2.4 Phytates

Phytic acid is a bioactive compound with wide distribution in plant foods. Due to its
molecular structure, phytic acid has affinity to polyvalent cations such as minerals
and trace elements. Phytic acid is one of the most important sources of phosphorus in
plant seeds, and sesame is no exception. In fact, sesame seeds are richer in phytate
than the commonly known legumes. In oil seeds such as sunflower, soybean,
sesame, linseed, and rapeseed, the phytic acid content ranges from 1% to 5.4%
compared to 0.2–2.9% in legumes. A defatted sesame meal has much higher phytate
concentration than that of soybean meal [98]. Graf and Dintzis [99] have measured
5.36% phytic acid content in sesame seeds. Often, phytates are termed as anti-
nutrient for preventing mineral absorption from meal, but it is also seen that phytates
have anticancerous and hypocholesterolemic activities [100, 101].

2.5 Polyunsaturated Fatty Acids

Fatty acids are carboxylic acids with long-chain hydrocarbon side groups derived
from or contained as esterified molecular form in lipids (fat, oil, or wax) of microbes,
animals, and plants usually ranging from 1 to 30 carbon atoms in length attached to a
terminal carboxyl group [102]. Polyunsaturated fatty acids (PUFAs) are long-chain
fatty acids containing two or more double bonds introduced by specific desaturase
enzymes. Over the years, vegetable oils rich in various PUFAs have emerged as
potential dietary elements for normal growth and development of human beings with
considerable biomedical significance. LC-x-3-PUFAs are important for human
health in maintaining the cellular membrane by regulating cholesterol synthesis,
transportation [103], and eicosanoid synthesis [104].
Sesame is a high-energy food containing approximately 50% oil. The fatty acid
composition of sesame oil is highly desirable with about 80–85% unsaturated acids
and only 15–20% of saturated acids. Sesame oil consists mainly of linoleic
Sesame: Bioactive Compounds and Health Benefits 11

(35–50%) and oleic (35–50%) acids, with small amount of palmitic (7–12%) and
stearic (3.5–6%) acids, but with only traces of linolenic acid [105, 106]. Studies
indicate that a high intake of n-6 fatty acids shifts the physiologic state to one that is
pro-thrombotic and pro-aggregatory, characterized by increase in blood viscosity,
vasospasm, and vasoconstriction and decrease in bleeding time. The n-3 fatty acids,
however, have anti-inflammatory, antithrombic, hypolipidemic, and vasodilatory
properties [103]. Sesame in combination with soybean oil increases the vitamin E
activity along with nutritive value of the lipid [19, 107].
Sesame varieties differ greatly in fatty acid contents of their seeds, and this under-
standing could lead to crop engineering with an aim to develop better quality edible oil in
future [108]. Indian sesame germplasms have lower level of saturated fatty acids
compared to other sesame varieties with palmitic acid being the prominent one. C18:1
and C18:2 are the major unsaturated fatty acids present in Indian varieties [109]. In
unsaturated fatty acids, while the content of C18:1 and C18:2 is quite high in sesame oil
(38–49% and 17–43%, respectively), the C18:3 content is below par (<1%) [110]. A
critical analysis of different cultivars is the need of hour, if we are assertive to improve the
nutritional quality of oil by engineering the fatty acid biosynthetic pathway.

2.6 Short-Chain Peptides, Protein Hydrolysates, and Their

Functional Properties

The bioactive polypeptides are amino acid chains joined by amide or peptide bonds,
with molecular weight not exceeding beyond 20 kDa. These are protein fragments
that have independent function in various biochemical, physiological, or cellular
processes. While some of these peptides are formed naturally, it is the artificial
protein hydrolysates that are much in use and are structurally and functionally a
diverse group of peptides. Proteins can be digested by proteases, or specific fragment
can be created in bio-fermenters utilizing microbes [111]. Sesame food preparation is
also an important source of protein [112]. Protein hydrolysates are widely in use as
nutritional supplement, functional ingredient, food flavor enhancer, pharmaceuticals,
and cosmetics [113, 114]. They display hormone- or drug-like activities and based
on the mode of action can be classified as antimicrobial, antithrombotic, antihyper-
tensive, opioid, immunomodulatory, mineral binding, and antioxidative [115]. Use
of papain for producing sesame protein hydrolysates having better functional prop-
erties, high storability, and emulsifying properties than that of the original sesame
protein lysate has been studied by Bandyopadhyay and Ghosh [116].

3 Quantitative Insight in Lignan and Tocopherol Content:

Interspecific and Intraspecific Variation

Sesame crops are of high nutritional value owing to the presence of antioxidants
lignans (sesamin, sesamolin, and sesamol) and tocopherols (α-, γ-, and δ-forms).
Separation of tocopherol homologues in sesame by RPLC on triacontyl (C30)
12 N. Pathak et al.

stationary phase has been successfully optimized. The reproducibility of the procedure
is in the range 1.7–3.9%, and recovery ranged from 91% to 99% [117]. Lignans and
tocopherol homologue contents in a wide collection of 143 sesame lines (wild species,
landraces, introgressed lines, and cultivars) collected from diverse agroecological
zones of India have been determined by reverse-phase HPLC (RP-HPLC) [118].
Screening of sesame germplasm revealed an exploitable level of variation in major
bioactive compounds. High levels of sesamin and γ-tocopherol suggested for the
efficient introduction of these lines in the trait enhancement of other oilseed crops.

4 Biotechnological Approaches for Sesame

Sesame crop is relatively superior in oil quantity and quality compared to many
major oil crops. The oil content varies from 35% to 60%, but mostly it is about 50%
of seed weight [119]. Therefore, genetic engineering of sesame directed toward
creation of sesame oil having diverse composition of lignans and tocopherols with
high nutritional value is an important biotechnological aspect. With high genetic
diversity, sesame becomes a promising oilseed crop for performing genetic manip-
ulations to obtain high yield and quality oil [120].

4.1 Genome Sequencing of Sesame Crop: Unraveling the Oil

Biosynthetic Pathway

Higher oil content compared to other oilseed crops, combined with small genome
related to oil quality enhancement, makes sesame an invaluable model plant for
studying oil biosynthesis. Wang et al. [121] have done de novo assembling of the
genome, where a set of 12 transcriptomes and 29 resequenced accessions provided
a large resource for exploring the mechanisms underlying different oil content
between sesame and soybean, as well as among the sesame accessions. The results
reveal the presence of whole genome duplication and absence of the Toll/interleukin-
1 receptor domain in resistance genes. Genes and oil biosynthetic pathways respon-
sible for high oil content were determined employing comparative genomic and
transcriptomic analyses. That has indicated for the expansion of type 1 lipid transfer
genes by tandem duplication, the contraction of lipid degradation genes, and the
differential expression of essential genes in the triacylglycerol biosynthesis pathway
in the early stage of seed development [121].
Resequencing data study in 29 sesame accessions from 12 countries suggests that
the high genetic diversity of lipid-related genes is linked with the wide variation in
oil content. Additionally, the results shed light on the pivotal stage of seed develop-
ment, oil accumulation, and potential key genes for sesamin production, an impor-
tant lignan with numerous health benefits. Recently, two more sesame landraces and
the chloroplast genome have been sequenced [121, 122] enriching the sesame
genome dataset. A recent genome-wide association study has identified SiNST1as
the candidate gene for lignin and cellulose biosynthesis, and this gene could
Sesame: Bioactive Compounds and Health Benefits 13

indirectly be associated with sesamin and sesamolin content [123]. Thankfully, a few
open access platform like Sinbase is also being maintained where all these datasets
are managed for further analysis and utilization by sesame researchers [121].

4.2 Functional Gene Expression: “Lignan Biosynthetic Genes”

High-throughput sequencing technology has provided ample data on DNA

sequences for the genomes of many plant species. Expressed sequence tags (EST)
of many crop species have been generated, and thousands of sequences have been
annotated as putative functional genes using powerful bioinformatics tools. Impact
on breeding programs could be reached with this approach, because quality of crop
plants is a direct function of their metabolite content [124] and quality of plant
tissues determines their commercial value with respect to flavor, fragrance, shelf life,
physical attributes, etc. [125].
Comparative analysis of expressed sequence tags from Sesamum indicum and
Arabidopsis thaliana developing seeds has been done by Suh et al. [126]. The group
could identify the genes involved in accumulation of seed storage products and in the
biosynthesis of lignans, sesamin, and sesamolin. Their study could also identify the
identical and different gene expression profiles during sesame and Arabidopsis seed
development and the genes specific to sesame seeds [126]. Sirato-Yasumoto et al.
[67] reported that sesamin content in sesame seeds was controlled by polygenes,
because F2 populations originating from reciprocal crosses between high sesamin
and normal sesamin varieties showed a continuous distribution in sesamin content,
and correlation coefficients between F2 and F3 generations were positive and highly
significant for sesamin content.
Ono et al. [29] reported that CYP81Q1 is a single gene encoding (þ)-piperitol/
(þ)-sesamin synthase in the Sesamum indicum genome, suggesting that CYP81Q1 is
a single enzyme, which catalyzes the formation of (þ)-sesamin from (þ)-
pinoresinol. The CYP81Q1 homologue (CYP81Q3) showed no activity of (þ)-
sesamin biosynthesis, due to (þ)-sesamin deficiency in this species, whereas S.
radiatum showed dual activity of the enzyme. Further, expression profile of
CYP81Q1 gene was temporally consistent with the accumulation pattern of (þ)-
sesamin during seed development. Hata et al. [127] detected sesamin (using ultra-
performance liquid chromatography-fluorescence detection) in sesame leaves of two
Japanese sesame varieties “Gomazou” and “Kin-goma,” which differed in sesamin
content of the seed, and probed genotypic differences. The higher sesamin content of
“Gomazou” leaves correlated well with that of seeds and the expression of the
sesamin biosynthetic gene CYP81Q1, indicating that genotypic difference of
CYP81Q1 gene expression affected leaf sesamin contents.
To comprehend sesame domestication, we examined the expression of sesamin
synthase (CYP81Q1) during capsule maturation in three wild Sesamum spp. and four
sesame cultivars [128]. Among the cultivars, only S. indicum (CO-1) exhibited
transcript abundance of sesamin synthase and high sesamin content similar to S.
14 N. Pathak et al.

malabaricum, whereas other cultivars had low expression, indicating that sesamin
synthase was not favored during domestication.

5 Conclusions

Sesame is no more an “orphan” crop – the widespread collection of varieties and

landraces coupled with extensive research in every aspect has made the crop as the
perfect model system amidst other oilseeds. The superior quality of sesame seed oil
containing a variety of lignans and tocochromanols merits a higher place among the
oilseed crops being consumed worldwide. The diversity of sesame cultivars and their
characterization has given sesame researchers enough impetus to create genetically
engineered sesame varieties with high yield of secondary metabolites. Sesame seeds
are microcapsules for health promotion and disease prevention in humans and an
sustained effort in this area of oilseed research would be of immense value to the
plant breeders as well as consumers.

Acknowledgments Ashwani K Rai gratefully acknowledges the National Academy of Sciences,

India, for awarding NASI-Senior Scientist Platinum Jubilee Fellowship. Niti Pathak wishes to thank
Dr. K V Bhat, NBPGR for the work carried out in his lab.

References
1. Ashri A (2007) Sesame (Sesamum indium L.) In: Singh RJ (ed) Genetic resources, chromo-
some engineering, and crop improvement, Oilseed crops, vol 4. CRC Press, Boca Raton,
pp 231–289
2. Joshi AB (1961) Sesamum. Indian Central Oilseed Committee, Hyderabad, pp 1–109
3. Weiss EA (1971) Sesame, castor and safflower, barnes and noble, World crop series. Leonard
Hill, New York, pp 311–525
4. Bedigian D, Seihler DS, Harlan JR (1985) Sesamin, sesamolin and the origin of sesame.
Biochem Syst Ecol 13:133–139
5. Bedigian D, Harlan JR (1986) Evidence for cultivation of sesame in the ancient world. Econ
Bot 40:137–154
6. USDA (2015) USDA national nutrient database for standard reference, release 18. U.S.
Department of Agriculture, Agricultural Research Service, Nutrient Data Laboratory,
Beltsville. http://www.nal.usda.gov/fnic/foodcomp
7. Pathak N, Rai AK, Ratna K, Bhat KV (2014) Value addition in sesame: a perspective on
bioactive components for enhancing utility and profitability. Pharmacogn Rev 8(16):147–155.
https://doi.org/10.4103/0973-7847.134249
8. Dimitrios B (2006) Sources of natural phenol antioxidants. Trends Food Sci Technol
17:505–512
9. Manach C, Williamson G, Morand C, Scalbert A, Remesy C (2005) Bioavailability and
bioefficacy of polyphenols in humans I- review of 97 bioavailability studies. Am J Clin Nutr
81:230S–242S
10. Temple NJ (2000) Antioxidants and disease: more questions than answers. Nutr Res
20:449–559
Sesame: Bioactive Compounds and Health Benefits 15

11. Kamal-Eldin A, Appelquist LA, Yousif G (1994) Lignan analysis in seed oils from four
sesamum species: comparison of different chromatographic methods. J Am Oil Chem Soc
71:141–145
12. Robinson R (1927) The relationship of some complex natural products to the simple sugars
and amino acids. Durham Univ Philos Soc 8:14–59
13. Haworth RD (1936) Natural resins. Annu Rep Progr Chem 33:266–279
14. Katsuzaki H, Osawa T, Kawakishi S (1994) Chemistry and antioxidative activity of lignan
glucosides in sesame seed. ACS Symp Ser 574:275–280
15. Katsuzaki H, Osawa T, Kawashiki S (1994) Chemistry and antioxidative activity of lignan
glucosides in sesame seed, Chapter 28. In: Food phytochemicals for cancer prevention, ACS
symposium series, vol 547. American Chemical Society, Washington, DC, pp 275–280
16. Moazzami AA, Andersson RE, Kamal-Eldin A (2006) HPLC analysis of sesaminol glucosides
in sesame seeds. J Agric Food Chem 54:633–638. https://doi.org/10.1021/jf051541g
17. Brar G, Ahuja KL (1979) Sesame: its culture, genetics, breeding and biochemistry. Annu Rev
Plant Sci 1:245–313
18. Yamashita K, Iizuka Y, Imai T, Namiki M (1995) Sesame seed and its lignans produce marked
enhancement of vitamin E activity in rats fed a low alpha- tocopherol diet. Lipids 30:
1019–1028
19. Namiki M (1995) The chemistry and physiological functions of sesame. Food Rev Int 11:
281–329
20. Kamal-Eldin A (2005) Minor components in vegetable oils. In: Shahidi F (ed) Baileys
industrial fats and oils. Chapter 12, edible oil and fat products: speciality oils and oil products.
Wiley, Sussex
21. Haller HL, Mc Govran ER, Goodhue LD, Sullivan WN (1942) The synergistic action of
sesamin with pyrethrum insecticides. J Org Chem 7(2):183–184
22. Jones WA, Beroza M, Decker ED (1962) Isolation and structure of sesangolin: a constituent of
Sesamum angolense. J Org Chem 27:3232–3235
23. Cassida JE, Engel JL, Essac EG, Kamienski FX, Kuwatsuka S (1966) Methylene-14C-
dioxyphenyl compounds: metabolism in relation to their synergistic action. Science 153:
1130–1133
24. Mathews CK, Van Holde KE, Ahern KG (2000) Biochemistry, 3rd edn, Benjamin/Cummings,
an imprint of Addison Wesley Longman, pp 700–704
25. Jain SC, Khanna P (1973) Production of sterols from Sesamum indicum L. tissue culture.
Indian J Pharm 35:163–164
26. Kato MJ, Chu A, Davin LB, Lewis NG (1998) Biosynthesis of antioxidant lignans in Sesamum
indicum seeds. Phytochemistry 47(4):583–591
27. Davin LB, Wang HB, Crowell AL, Bedgar DL, Martin DM, Sarkanen S, Lewis NG (1997)
Stereoselective bimolecular phenoxy radical coupling by an auxiliary (dirigent) protein with-
out an active center. Science 275:362–366
28. Jiao Y, Davin LB, Lewis NG (1998) Furanofuran lignan metabolism as a function of seed
maturation in Sesamum indicum: methylenedioxy bridge formation. Phytochemistry 49:
387–394
29. Ono E, Nakai M, Fukui Y, Tomimori N, Fukuchi-Mizutani M, Saito M, Satake H, Tanaka T,
Katsuta M, Umezawa T, Tanaka Y (2006) Formation of two methylenedioxy bridges by a
Sesamum CYP81Q protein yielding a furofuran lignan, (þ)-sesamin. Proc Natl Acad Sci USA
103(26):10116–10121
30. Yoshida Y, Niki E, Noguchi N (2003) Comparative study on the action of tocopherols and
tocotrienols as antioxidant: chemical and physical effects. Chem Phys Lipids 123(1):63–75
31. Hofius D, Sonnewald U (2003) Vitamin E biosynthesis: biochemistry meets cell biology.
Trends Plant Sci 8(1):6–8
32. Brigelius-Flohe R, Traber MG (1999) Vitamin E: function and metabolism. FASEB J 13(10):
1145–1155
16 N. Pathak et al.

33. Colombo ML (2010) An update on vitamin E, tocopherol and tocotrienol- perspectives.

Molecules 15(4):2103–2113. https://doi.org/10.3390/molecules15042103
34. Bramley PM, Elmadfa I, Kafatos A, Kelly FJ, Manios Y, Rexborough HE, Schuch W,
Sheehy PJA, Wagner KH (2000) Vitamin E. J Sci Food Agric 80:913–938
35. Herbers K (2003) Vitamin production in transgenic plants. J Plant Physiol 160:821–829.
https://doi.org/10.1078/0176-1617-01024
36. Franzen JJ, Bausch D, Glatze D, Wagner E (1991) Distribution of vitamin E in spruce seedling
and mature tree organs, and within the genus. Phytochemistry 30:147–151
37. Hassapidou MN, Manoukas AG (1993) Tocopherol and tocotrienol compositions of raw table
olive fruit. J Sci Food Agric 61(2):277–280
38. DellaPenna (2005) Progress in the dissection and manipulation of vitamin E synthesis. Trends
Plant Sci 10:574–579. https://doi.org/10.1016/j.tplants.2005.10.007
39. Norris SR, Shen X, DellaPenna D (1998) Complementation of the Arabidopsis pds1 mutation
with the gene encoding p-hydroxyphenylpyruvate dioxygenase. Plant Physiol 117:1317–1323
40. Collakova E, DellaPenna D (2001) Isolation and functional analysis of homogentisate
phytyltransferase from Synechocystis sp. PCC 6803 and Arabidopsis. Plant Physiol 127:
1113–1124
41. Savidge B, Weiss JD, Wong YHH, Lassner MW, Mitsky TA, Shewmaker CK, Beittenmiller D,
Valentin HE (2002) Isolation and characterization of homogentisate phytyltransferase genes
from Synechocystis sp. PCC 6803 and Arabidopsis. Plant Physiol 129:321–322
42. Cheng Z, Sattler S, Maeda H, Sakuragi Y, Bryant DA, Dellapenna D (2003) Highly divergent
methyltransferases catalyze a conserved reaction in tocopherol and plastoquinone synthesis in
cyanobacteria and photosynthetic eukaryotes. Plant Cell 15:2343–2356
43. Van Eenennaam AL, Lincoln K, Durett TP, Valentin HE, Shewmaker CK, Thorne GM, Jiang J,
Baszis SR, Levering CK, Aasen ED, Hao M, Stein JC (2003) Engineering vitamin E content:
from Arabidopsis mutant to soy oil. Plant Cell 15(12):3007–3019
44. Porfirova S, Bergmüller E, Tropf S, Lemke R, Dörmann P (2002) Isolation of an Arabidopsis
mutant lacking vitamin E and identification of a cyclase essential for all tocopherol biosyn-
thesis. Proc Natl Acad Sci U S A 99:12495–12500
45. Sattler SE, Cajon EB, Coughlin SJ, DellaPenna D (2003) Characterization of tocopherol
cyclases from higher plants and cyanobacteria: evolutionary implications for tocopherol
synthesis and function. Plant Physiol 132:2184–2195
46. Budowski P, Markley KS (1951) The chemical and physiological properties of sesame oil.
Chem Rev 48:125–151
47. Osawa T, Nagata M, Namiki M, Fukuda Y (1985) Sesamolinol, a novel antioxidant isolated
from sesame seeds. Agric Biol Chem 49:3351–3352
48. Hirata F, Fujita K, Ishikura Y, Hosoda K, Ishikawa T, Nakamura H (1996) Hypercholesterol-
emic effect of sesame lignan in human. Atherosclerosis 122:135–136
49. Shimizu S, Akimoto K, Shinmen Y, Kawashima H, Sugano M, Yamada H (1991) Sesamin is a
potent and specific inhibitor of delta-5-desaturase in polyunsaturated fatty acid biosynthesis.
Lipids 26:512–516
50. Hirose N, Inoue T, Nishihara K, Sugano M, Akimoto K, Shimizu S, Yamada S (1991)
Inhibition of cholesterol absorption and synthesis in rats by sesamin. J Lipid Res 32:629–638
51. Yokota T, Matsuzaki Y, Koyama M, Hitomi T, Kawanaka M, Enoki-Konish M, Okuyama Y,
Takayasu J, Nishino H, Nishikawa A, Osawa T, Sakai T (2007) Sesamin, a lignan of sesame,
down-regulates cyclin D1 protein expression in human tumor cells. Cancer Sci 98(9):
1447–1453. https://doi.org/10.1111/j.1349-7006.2007.00560.x
52. Hsu DZ (2005) Effect of sesame oil on oxidative-stress-associated renal injury in endotoxemic
rats: involvement of nitric oxide and proinflammatory cytokines. Shock 24:276–280
53. Ashakumary L, Rouyer I, Takahashi Y, Ide T, Fukuda N, Aoyama T, Hashimoto T, Mizugaki
M, Sugano M (1999) Sesamin, a sesame lignan, is a potent inducer of hepatic fatty acid
oxidation in the rat. Metabolism 48:1303–1313
Sesame: Bioactive Compounds and Health Benefits 17

54. Nonaka M, Yamashita K, Izuka Y, Namiki M (1997) Effects of sesaminol and sesamin on
eicosanoid production and immunoglobulin level in rats given ethanol. Biosci Biotechnol
Biochem 61:836–839
55. Lee CC, Chen PR, Lin S, Tsai SC, Wang BW, Chen WW (2004) Sesamin induces nitric oxide
and decreases endothelin-1 production in HUVECs: possible implications for its antihyper-
tensive effect. J Hypertens 22:2329–2338
56. Nakano D, Kurumazuka D, Nagai Y, Nishiyama A, Kiso Y, Matsumura Y (2008) Dietary
sesamin suppresses aortic NADPH oxidase in DOCA salt hypertensive rats. Clin Exp
Pharmacol Physiol 35(3):324–326. https://doi.org/10.1111/j.1440-1681.2007.04817.x
57. Cheng FC, Jinn TR, Hou RC, Tzen JTC (2006) Neuroprotective effects of sesamin and
sesamolin on gerbil brain in cerebral ischemia. Int J Biomed Sci 2(3):284–288
58. Hemalatha S, Ghafoorunissa (2004) Lignans and tocopherols in Indian sesame cultivars. J Am
Oil Chem Soc 81:467–470
59. Abe C, Ikeda S, Yamashina K (2005) Dietary sesame seeds elevate α-tocopherol concentration
in rat brain. J Nutr Sci Vitaminol 51:223–230
60. Kamal-Eldin A, Pettersson D, Appelqvist LÅ (1995) Sesamin (a compound from sesame oil)
increases tocopherol levels in rats fed ad libitum. Lipids 30:499–505
61. Wu WH, Kang YP, Wang NH, Jou HJ, Wang TA (2006) Sesame ingestion affects sex
hormones, antioxidant status, and blood lipids in postmenopausal women. J Nutr 136(5):
1270–1275
62. Mak DHF, Po YC, Kam MK (2011) Antioxidant and anti-carcinogenic potentials of sesame
lignans. In: Bedigian D (ed) Sesame the genus sesamum. CRC Press, Boca Raton
63. Sandra MS, Lilian UT (2011) Sesame seeds and its lignans: metabolism and bioactivities. In:
Bedigian D (ed) Sesame the genus Sesamum. CRC Press, Boca Raton
64. Matsumara Y, Kita S, Tanida Y, Taguchi S, Morimoto S, Akimoto K, Tanaka T (1998)
Antihypertensive effect of sesamin, protection against development and maintenance of
hypertension in stroke-prone spontaneously hypertensive rats. Biol Pharm Bull 21:469–473
65. Chavali SR, Zhong WW, Forse RA (1998) Dietary α-linolenic acid increases TNF-α, and
decreases IL-6, IL-10 in response to LPS: effect of sesamin on the Δ-5 desaturation of ω6 and
ω3 fatty acids in mice. Prostaglandins Leukot Essent Fat Acids 58(3):185–191
66. Lim JS, Adachi Y, Takahashi Y, Ide T (2007) Comparative analysis of sesame lignans (sesamin
and sesamolin) in affecting hepatic fatty acid metabolism in rats. Br J Nutr 97(1):85–95.
https://doi.org/10.1017/S0007114507252699
67. Sirato-Yasumoto S, Katsuta M, Okuyama Y, Takahashi Y, Ide T (2001) Effect of sesame seeds
rich in sesamin and sesamolin on fatty acid oxidation in rat liver. J Agric Food Chem
49:2647–2651
68. Hirose N, Doi F, Ueki T, Akazawa K, Chijiiwa K (1992) Suppressive effect of sesamin against
7, 12-dimethylbenz[a]-anthracene induced rat mammary carcinogenesis. Anticancer Res
12:1259–1265
69. Coulman KD, Liu Z, Quan HW, Michaelides J, Thompson LU (2005) Whole sesame seed is as
rich a source of mammalian lignan precursors as whole flaxseed. Nutr Cancer 52:156–165.
https://doi.org/10.1207/s15327914nc5202_6
70. Liu Z, Saarinen NM, Thompson LU (2006) Sesamin is one of the major precursors of
mammalian lignans in sesame seed (Sesamum indicum) as observed in vitro and in rats.
J Nutr 136:906–912
71. Penalvo JL, Heinonen SM, Aura AM, Adlercreutz H (2005) Dietary sesamin is converted to
enterolactone in humans. J Nutr 135:1056–1062
72. Annussek G (2001) Sesame oil in: gale encyclopedia of alternative medicine. Gale Group and
Looksmart, Detroit
73. Ang ES, Lee ST, Gan CS, See PG, Chan YH, Nag LH, Machin D (2001) Evaluating the role of
alternative therapy in burn wound management: randomized trial comparing moist exposed
burn ointment with conventional methods in the management of patients with second- degree
burns. Med Gen Med 3:2–7
18 N. Pathak et al.

74. Yong YL (1999) Analysis of MEBO cream, Report no. 99033191. Institute of Science and
Forensic Medicine, Department of Scientific Services, Health Science Division, Singapore
75. Kamal-Eldin A, Appelqvist LÅ (1996) The chemistry and antioxidant properties of tocoph-
erols and tocotrienols. Lipids 31:671–701
76. Burton GW, Traber MG (1990) Vitamin E in antioxidant activity biokinetics and bioavailabil-
ity. Annu Rev Nutr 10:375–382. https://doi.org/10.1146/annurev.nu.10.070190.002041
77. Burton GW (1994) Vitamin E: molecular and biological function. Proc Nutr Soc 53(2):
251–262
78. Li D, Saldeen T, Romeo F, Mehta JL (1999) Relative effects of alpha- and gamma-tocopherol
on low-density lipoprotein oxidation and superoxide dismutase and nitric oxide synthase
activity and protein expression in rats. J Cardiovasc Pharmacol Ther 4:219–226
79. Saldeen T, Engström K, Jokela R, Wallin R (1999) Natural antioxidants and anticarcinogens in
nutrition, health and disease. In: Importance of in vitro stability for in vivo effects of fish oils.
The Royal Society of Chemistry, Cambridge, UK, Special Publication 240, pp 326–330
80. Qureshi AA, Bradlow BA, Brace L, Manganello J, Peterson DM, Pearce BC, Wright JJK,
Gapor A, Elson CE (1995) Response of hypercholesterolemic subjects to administration of
tocotrienols. Lipids 30(12):1171–1177
81. Schwenke DC (2002) Does lack of tocopherols and tocotrienols put women at increased risk of
breast cancer? J Nutr Biochem 13(1):2–20
82. Olcott HS, Emerson OH (1937) Antioxidants and the autoxidation of fats, IX, the antioxidant
properties of the tocopherols. J Am Oil Chem Soc 59(6):1008–1009
83. Girotti AW (1998) Lipid hydroperoxide generation, turnover, and effector action in biological
systems. J Lipid Res 39:1529–1542
84. Liebler DC (1993) The role of metabolism in the antioxidant functions of vitamin E. Crit Rev
Toxicol 23:147–169. https://doi.org/10.3109/10408449309117115
85. Dabrowski KJ, Sosulski F (1984) Quantification of free and hydrolizable phenolic acids in
seeds by capillary gas liquid chromatography. J Agric Food Chem 32(1):123–127
86. Feroj-Hasan AFM, Begu S, Furumoto T, Fukui H (2000) A new chlorinated red napthaquinone
from roots of Sesamum indicum. Biosci Biotechnol Biochem 64:873–874. https://doi.org/10.1
271/bbb.64.873
87. Lyon CK (1972) Sesame, present knowledge of composition and use. J Am Oil Chem Soc
49:245–249
88. Shimoda T, Takabayashi J, Ashira W, Takafuji (1997) Response of predatory insect
Scolothrips takahashi towards herbivore induced plant volatiles under laboratory and field
conditions. J Chem Ecol 23:2033–2048
89. Salunkhe DK, Chavan JK, Adsule RN, Kadam SS (1991) World oilseeds: chemistry, technol-
ogy and utilization. Springer, New York, pp 1–554
90. Van Rensburg SJ, Daniels WM, Van Zyl JM, Taljaard JJ (2000) A comparative study of the
effects of cholesterol, beta-sitosterol, beta-sitosterol glucoside, dehydroepiandrosterone sul-
phate and melatonin on in vitro lipid peroxidation. Metab Brain Dis 15:257–265
91. Bouic PJ (2002) Sterols and sterolins: new drugs for the immune system? Drug Discov Today
7:775–778
92. Zhao W, Miao X, Jia S, Pan Y, Huang Y (2005) Isolation and characterization of microsatellite
loci from the mulberry Morus L. Plant Sci 168:519–525
93. Moreau RA, Whitaker BD, Hicks Kevin B (2002) Phytosterols, phytostanols, and their
conjugates in foods: structural diversity, quantitative analysis, and health-promoting uses.
Prog Lipid Res 41:457–500
94. Mohamed HM, Awatif II (1998) The use of sesame oil unsaponifiable matter as a natural
antioxidant. Food Chem 62:269–276
95. Gharby S, Harhar H, Bouzoubaa Z, Asdadi A, El Yadini A, Charrouf Z (2015) Chemical
characterization and oxidative stability of seed and oil of sesame grown in Morocco. J Saudi
Soc Agric Sci 16:105–111. https://doi.org/10.1016/j.jssas.2015.03.004
Sesame: Bioactive Compounds and Health Benefits 19

96. Pegel KH (1997) The importance of sitosterol and sitosterolin in human and animal nutrition.
S Afr J Sci 93:263–268
97. Nieman DC (1994) Exercise, infection and immunity. Int J Sports Med 15:131–141. https://
doi.org/10.1055/s-2007-1021128
98. de Boland AR, Garner GB, O’Dell BL (1975) Identification and properties of “phytate” in
cereal grains and oilseed products. J Agric Food Chem 23:1186–1189
99. Graf E, Dintzis FR (1982) High-performance liquid chromatographic method for the determi-
nation of phytate. Anal Biochem 119:413–417
100. Urbano G, López-Jurado M, Aranda P, Vidal-Valverde C, Tenorio E, Porres J (2000) The role
of phytic acid in legumes: antinutrient or beneficial function? J Physiol Biochem 56:283–294
101. Kuroda Y, Shamsuddin AM (1995) Inositol phosphates have novel anticancer function. J Nutr
125:725S–732S
102. Gunstone F, Harwood JL, Padley FB (1994) The lipid handbook, 2nd edn. Chapman and Hall,
London, pp 47–208
103. Simopoulos AP (1999) Essential fatty acids in health and chronic disease. Am J Clin Nutr 70(3
Suppl):560S–569S
104. Kankaanpaa P, Sutas Y, Salminen S, Lichtenstein A, Isolauri E (1999) Dietary fatty acids and
allergy. Ann Med 31:282–287
105. Kamal-Eldin A, Appelqvist LÅ (1994) Variation in the composition of sterols, tocopherols and
lignans in seed oils from four Sesamum species. J Am Oil Chem Soc 71:149–156
106. Spencer GF, Herb SF, Gormisky PJ (1976) Fatty acid composition as a basis for identification
of commercial fats and oils. J Am Oil Chem Soc 53:94–96
107. Shahidi F, Tan Z (2011) Physiological effects of sesame bioactive and antioxidant compounds.
In: Bedigian D (ed) Sesame the genus sesamum. CRC Press, Boca Raton
108. Uzun B, Arslan C, Furat S (2008) Variation in fatty acid compositions, oil content and oil yield
in germplasm collection of sesame (Sesamum indicum L.) J Am Oil Chem Soc 85:1135–1142
109. Mondal N, Bhat KV, Srivastava PS (2010) Variation in fatty acid composition in Indian
germplasm of sesame. J Am Oil Chem Soc 87(11):1263–1269
110. Bhunia RK, Chakraborty A, Kaur R et al (2015) Analysis of fatty acid and lignan composition
of Indian germplasm of sesame in terms of their nutritional merits. J Am Oil Chem Soc 92:
65–76
111. Aluko R (2012) Bioactive peptides. In: Functional foods and nutraceuticals, Food science text
series. Springer, New York, pp 37–61
112. Dench JE, Rivas N, Caygill JC (1981) Selected functional properties of sesame (Sesamum
indicum L.). Flour and two protein isolates. J Sci Food Agric 32:557–564. https://doi.org/
10.1002/jsfa.2740320606
113. Frokjaer S (1994) Use of hydrolysates for protein supplementation. Food Technol 48:86–88
114. Giese J (1994) Proteins as ingredients: types, functions, applications. Food Technol 48:50–60
115. Sánchez A, Vázquez A (2017) Bioactive peptides: a review. Food Qual Saf 1(1):29–46. https://
doi.org/10.1093/fqs/fyx006
116. Bandyopadhyay K, Ghosh S (2002) Preparation and characterization of papain-modified
sesame (Sesamum indicum L.) protein isolates. J Agric Food Chem 50(23):6854–6857
117. Saha S, Walia S, Kundu A, Pathak N (2013) Effect of mobile phase on resolution of the
isomers and homologues of tocopherols on a triacontyl stationary phase. Anal Bioanal Chem
405:9285–9295. https://doi.org/10.1007/s00216-013-7336-9
118. Pathak N, Rai AK, Saha S, Walia SK, Sen SK, Bhat KV (2014) Quantitative dissection of
antioxidative bioactive components in cultivated and wild sesame germplasm reveals poten-
tially exploitable wide genetic variability. J Crop Sci Biotechnol 17(3):127–139
119. Ashri A, Downey RK, Robbelen G (1989) Brassica species. In: Ashri A, Robbelen G, Downey
RK (eds) Oil crops of the world. McGraw-Hill, New York, pp 339–382
120. Pathak N, Rai AK, Kumari R, Thapa A, Bhat KV (2014) Sesame crop: an underexploited
oilseed holds tremendous potential for enhanced food value. Agric Sci 5(6):519–529. https://
doi.org/10.4236/as.2014.56054
 20 N. Pathak et al.

121. Wang L, Yu S, Tong C, Zhao Y, Liu Y, Song C, Zhang Y, Zhang X, Wang Y, Hua W, Li D, Li
D, Li F, Yu J, Xu C, Han X, Huang S, Tai S, Wang J, Xu X, Li Y, Liu S, Varshney RK, Wang J,
Zhang X (2014) Genome sequencing of the high oil crop sesame provides insight into oil
biosynthesis. Genome Biol 15:R39. https://doi.org/10.1186/gb-2014-15-2-r39
122. Wei X, Zhu X, Yu J, Wang L, Zhang Y, Li D, Zhou R, Zhang X (2016) Identification of sesame
genomic variations from genome comparison of landrace and variety. Front Plant Sci 7:1169.
https://doi.org/10.3389/fpls.2016.01169
123. Wei X et al (2015) Genetic discovery for oil production and quality in sesame. Nat Commun
6:8609. https://doi.org/10.1038/ncomms9609
124. Memelink J (2004) Tailoring the plant metabolome without a loose stitch. Trends Plant Sci
7:305–307. https://doi.org/10.1016/j.tplants.2005.05.006
125. Hall C, Tulbek MC, Xu Y (2006) Flaxseed. Adv Food Nutr Res 51:1–97
126. Suh MC, Kim MJ, Hur CG, Bae JM, Park YI, Chung CH, Kang CW, Ohlrogge JB (2003)
Comparative analysis of expressed sequence tags from Sesamum indicum and Arabidopsis
thaliana developing seeds. Plant Mol Biol 52(6):1107–1123
127. Hata N, Hayashi Y, Okazawa A, Ono E, Satake H, Kobayashi A (2010) Comparison of
sesamin contents and CYP81Q1 gene expressions in aboveground vegetative organs between
two Japanese sesame (Sesamum indicum L.) varieties differing in seed sesamin contents. Plant
Sci 178(6):510–516
128. Pathak N, Bhaduri A, Bhat KV, Rai AK (2015) Tracking sesamin synthase gene expression
through seed maturity in wild and cultivated sesame species – a domestication footprint. Plant
Biol 17(5):1039–1046. https://doi.org/10.1111/plb.12327

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