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Critical Reviews in Food Science and Nutrition
ISSN: 1040-8398 (Print) 1549-7852 (Online) Journal homepage: http://www.tandfonline.com/loi/bfsn20
Nutritional chemistry of the peanut (Arachis
hypogaea)
Ondulla T. Toomer
To cite this article: Ondulla T. Toomer (2017): Nutritional chemistry of the peanut (Arachis
hypogaea), Critical Reviews in Food Science and Nutrition, DOI: 10.1080/10408398.2017.1339015
To link to this article: https://doi.org/10.1080/10408398.2017.1339015
Accepted author version posted online: 29
Jun 2017.
Published online: 11 Oct 2017.
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Nutritional chemistry of the peanut (Arachis hypogaea)
Ondulla T. Toomer
United States Department of Agriculture-Agricultural Research Service, Market Quality and Handling Research Unit, Raleigh, NC, USA
ABSTRACT
Peanuts, Arachis hypogaea, are one of the most widely consumed legumes globally due to its nutrition, taste,
and affordability. Peanuts are protein and energy-rich and have been utilized worldwide to address the
nutritional needs in developing countries. Currently, its role in a heart-healthy diet has warranted
tremendous attention among consumer groups and within the scientific community. Additionally, current
studies have identified the value in the phytonutrient composition of peanuts, such as resveratrol,
isoflavonoids, phenolic acids, and phytosterols, which may enhance overall health and wellness. This article
presents a comprehensive review of the nutritional chemistry of peanut components (macronutrients—
proteins, lipids, carbohydrates; micronutrients—vitamins, minerals, phytonutrients) as related to health and
use within the body. An improved comprehensive knowledge and better understanding of the nutritional
chemistry of peanuts enables us to better harness the power of these nutrients in improved peanut products
within the food and feed industry.
KEYWORDS
Peanut nutrition; peanut
nutritional chemistry;
phytonutrients
Introduction
Peanut seeds are approximately 22 to 30% crude protein (Patee
and Young, 1982; Settaluri et al., 2012) and are a great vegetar-
ian source of protein and healthy fats. However, in years past
peanuts and tree nuts were perceived as an unhealthy food due
to their high fat content of 50% w/w (Zhao et al., 2012). Cur-
rent research studies have demonstrated that dietary inclusion
of peanuts and tree nuts has been linked to reduced heart dis-
ease (Jones et al., 2014), certain types of cancers (Gonzalez and
Salas-Salvad
o, 2006), and improved weight management
(Moreno et al., 2013). A study published in the New England
Journal of Medicine reported that eating nuts daily can reduce
death from heart disease by 29%, and even eating peanuts just
twice a week can reduce risk by 24% (Boa et al., 2013). Other
studies have demonstrated that regular peanut consumption
helps to decrease blood pressure among hypertensive individu-
als with significant reduction in diastolic blood pressure (Jones
et al., 2014).
In parallel, four large epidemiological studies (Adventist
Health Study, Iowa Women’s Health Study, Nurses’Health
Study, and Physician’s Health Study) examined the effect of fre-
quent nut consumption (tree nuts and peanuts) on the risk of
coronary and ischemic heart disease. The Adventist Health
Study (Fraser et al., 1992) demonstrated that subjects who con-
sumed nuts more than four times a weeks experienced signifi-
cantly fewer coronary heart disease-related events and fewer
definite fatal coronary heart disease-related events (Francisco
and Resurreccion, 2008). The Nurses’Health Study showed
that women who consumed more than one ounce of nuts per
week had significantly lower risk of total coronary heart disease
in comparison to women who consumed less than one ounce
of nuts per month (Hu et al., 1998). Similarly, the Iowa
Women’s Health Study demonstrated that regular consump-
tion of nuts significantly reduced the risk of coronary heart dis-
ease-related deaths in postmenopausal women (Ellsworth et al.,
2001). In addition, the Physician’s Health Study (Albert et al.,
2002) reported that deaths related to cardiac events were signif-
icantly reduced in subjects who regularly consumed nuts
(tree nuts and peanuts). Hence, today the overall image and
consumer perception of peanuts has shifted from an energy
dense food to a beneficial food item associated with improved
health benefits.
The peanut also known as the groundnut and/or goober is a
legume and taxonomically classified as Arachis hypogaea,is
believed to have originated in Central and South America with
cultivation spread to other parts of the world (Settaluri et al.,
2012). Today, peanuts are cultivated in China, India, Africa,
Japan, South America, and the United States, with more than
300 varieties grown worldwide (Settaluri et al., 2012). The four
market types of peanuts grown commercially in the United
States are runner, Virginia, Spanish, and Valencia.
The runner cultivar predominates 80% of the United States
peanut market and is primarily used for peanut butter, and has
an attractive uniform kernel size (American Peanut Council,
2014). Runners are predominately grown in Georgia, Alabama,
Florida, Texas, and Oklahoma. The Virginia type peanuts have
the largest kernel size and processed-in-shell are grown mainly
in Virginia and North Carolina for gourmet snacks, and com-
prises approximately 15% of the United States peanut market
(American Peanut Council, 2014). Spanish peanuts have a
CONTACT Ondulla T. Toomer Ondulla.Toomer@ars.usda.gov United States Department of Agriculture-Agricultural Research Service, Market Quality and
Handling Research Unit, 400 Dan Allen Drive, Schaub Hall, Box 7624, Raleigh, NC 27695, USA.
Color versions of one or more of the figures in the article can be found online at www.tandfonline.com/bfsn.
© 2017 Taylor & Francis Group, LLC
CRITICAL REVIEWS IN FOOD SCIENCE AND NUTRITION
https://doi.org/10.1080/10408398.2017.1339015
smaller kernel size with a higher oil content than other peanut
types are often used in peanut candies and snacks. Spanish type
peanuts are grown in Oklahoma and Texas and compromise
approximately 4% of the United States peanut market
(American Peanut Council, 2014). Valencia type peanuts are
produced primarily in New Mexico and comprise approxi-
mately 1% of the US peanut market (American Peanut Council,
2014). Valencia type peanuts have three or more small kernels
to a pod and are very sweet in taste and are therefore are usu-
ally roasted and sold in-the-shell.
Peanuts can be eaten either raw, boiled, or roasted and are
widely used to prepare a variety of packaged foods (peanut but-
ter, candies, confections, and snack products) in the United
States and are relied upon as a protein extender in developing
countries.
This comprehensive review aims to focus on the nutritional
chemistry of peanut components as related to health and use
within the body. In contrast, previous peanut nutrition reviews
have focused primarily on peanut allergens (S
aiz et al., 2013)
and/or peanut phytonutrients exclusively (Francisco and Res-
urreccion, 2008; Sales and Resurreccion, 2014). All foods are
composed of chemical compounds, which can be defined as
macro- or micro-nutrients such as proteins, carbohydrates,
fats, or vitamins, minerals and phytonutrients, respectively.
Macronutrients
Proteins
As peanuts are technically legumes they are more closely
related to chickpeas and soybeans than to almonds, walnuts, or
other tree nuts, and are more protein rich, and more nutrition-
ally complete than tree nuts (Ros, 2010). Iqbal et al. (2016)
reported that peanut proteins sampled from peanut seeds from
different geographic locations around the world were the same,
however with differing amounts. The peanut seed contains 32
different proteins (Pele, 2010) with only 18 of these proteins
identified as allergenic (Yusnawan et al., 2012). To date, 17
peanut proteins (Ara h 1 through Ara h 17) have been identi-
fied as peanut allergens responsible for peanut allergy by the
World Health Organization and International Union of Immu-
nological Societies (WHO/IUIS, 2017) Allergen Nomenclature
Sub-Committee, with Ara h 1, Ara h 2, and Ara h 6 being major
peanut allergens (Porterfield et al., 2009). Nevertheless, only
Ara h 1 through Ara h 10 have been well identified and bio-
chemically characterized (Table 1). Other studies have defined
Ara h 2 and Ara 6 as the major peanut allergens due to aller-
genic potency in allergenic-mediated assays (Zhuang and
Dreskin, 2013). Interestingly, peanut allergens in particular
have been shown to be extremely resistant to proteolytic diges-
tion, and heat or chemical denaturation (Koppelman et al.,
2010; Iqbal and Ateeq, 2013, Toomer et al., 2013). Other
reports have demonstrated that allergenic food proteins are
stable to digestion, and a common feature of many food
allergens (Wickham et al., 2009).
These allergenic proteins found within peanut encompass
seven different protein families, separated into two major pro-
tein fractions albumins and globulins (Patee and Young, 1982;
Settaluri et al., 2012). The globulin fractions were first catego-
rized as storage or reserve proteins of the peanut seed by Johns
and Jones (1916) and by Jones and Horn (1930). The two-glob-
ulin fractions, arachin, and non-arachin compromise approxi-
mately 87% of the peanut seed proteins (Basha and Pancholy,
1981).
Globulin proteins
Ara ha 1 is a glycoprotein belonging to the vicilin (7S) legume
globulin family. It compromises approximately 12–16% of pea-
nut proteins (deJong et al., 1998) and affects 35–95% of the
peanut-allergic population (Mari et al., 2006). Native Ara h 1
exists as a trimer formed by three identical monomers, with its
basic structure and core region very similar to other 7S globu-
lins. Mature Ara h 1 has 21 linear epitopes, with 14 within the
core region of the protein. However, in the native trimer forma-
tion these epitopes are either slightly or significantly buried
Table 1. Proteins Found in Whole Conventional Peanuts.
Protein Superfamily Protein Family Allergen Isoforms MW (kDa)
Prevalence of
IgE Binding Biological Function
Cupins Globulin-Vicilin-Type (7 S globulin) Ara h 1 Ara h 1.0101 64 (trimer
180 kDa)
35-95%
Globulin-Glycinin-type (11 S globulin) Ara h 3 Ara h 3.0101, Ara h 3.0201 60 50% Trypsin Inhibitor
Globulin-Glycinin-type (11 S globulin) Ara h 4
Ara h 3.0201 60 >50%
Prolamin Albumins-Conglutin (2 S Albumin) Ara h 2 Ara h 2.0101, Ara h 2.0201 17 >95% Trypsin Inhibitor
Albumins-Conglutin (2 S Albumin) Ara h 6 Ara h 6.0101 15 38%
Albumins-Conglutin (2 S Albumin) Ara h 7 Ara h 7.0101, Ara h 7.0201 15 13%
<
Non-Specific Lipid Transfer Proteins Ara h 9 Ara h 9.0101, Ara h 9.0201 9.8 45% Transport between
cell membranes
Profilin Profilin Ara h 5 Ara h 5.0101 15 13%
<
Regulate
polymerization of
actin
Bet v 1 Family Pathogenesis-Related Protien Ara h 8 Ara h 8.0101, Ara h 8.0201 17 70% plant protection
pathogen invasion
Glycosyl transferase GT-C Oleosin Ara h 10 Ara h 10.0101, Ara h
10.0201
16 21% Structural stability in
plant oil bodies
Ara h 11 Ara h 11.0101 14 not known
Scorpion toxin-like knottin Defensins Ara h 12 Arah 12.0101 5 to 12 not known host defense peptides
Ara h 13 Ara h 13.0101 5 to 12 not known host defense peptides
Ara h 4 found to be an isomer of Ara h 3 Drenamed Ara h 3.02
<D5 patients out of 40 were IgE reactive
2O. T. TOOMER
explaining the relatively weak Immunoglobulin E (IgE) reactiv-
ity in the native form, and strong IgE reactivity in the
denatured form (Zhou et al., 2013).
Ara h 3 is a seed storage protein belonging to the legumin
(11S globulin) family (Koppelman et al., 2003)andisIgEimmu-
noreactive in approximately 50% of peanut allergic patients and
functions as a trypsin inhibitor (Dodo et al., 2004; Wen et al.,
2007).MatureArah3isahexamer(360–380 kDa) formed by
two trimers (Jin et al., 2009), with each monomer having four
linear epitopes (Rabjohn et al., 1999). Ara h 3 in the native form
has the fourth epitope fully exposed, while the other three epito-
pes are completely or almost buried; suggesting that epitopes 1
and 2 may not be recognized by IgE antibodies in the native
form, while epitopes 4 and part of epitope 3 may be reactive in
the native form of Ara h 3 (Jin et al., 2009). Ara h 4 is an
isoform of Ara h 3 (Table 1) and is no longer thought to be a
distinct allergen and thus has been renamed to Ara h
3.02 (WHO/IUIS Allergen Nomenclature Sub-Committee, 2017).
Albumin proteins
Ara h 2 is a glycoprotein and accounts for approximately 6 to
9% of total peanut protein (Koppelman et al., 2001)witha
molecular weight of approximately 17 kDa (S
aiz et al., 2013).
Ara h 2 is a 2S albumin also known as conglutin and functions
as a trypsin inhibitor (Table 1), proteins that reduce the biologi-
cal activity of trypsin the digestive enzyme responsible for dietary
protein digestion (Maleki et al., 2003). Over 95% of the peanut
allergy population in the United States are Ara h 2 IgE reactive
(Koppelman et al., 2004; Scurlock and Burks, 2004;Palmeretal.,
2005) and therefore the most potent peanut allergen among pea-
nut-sensitive subjects and therefore clinically important in peanut
allergen sensitivity. Structurally, Ara h 2 has five a-helices
arranged in right-handed super helix connected by several
extended loops with four conserved disulfide bridges and 10
highly exposed epitope binding sites (Zhou et al., 2013).
Ara h 6, also a conglutin (2S albumin) protein, has a molec-
ular weight of 15 kDa and is 59% homologous to Ara h 2 with
similar allergenicity (Koppelman et al., 2005; Chen et al., 2013).
Studies by Suhr et al. (2004) and Lehmann et al. (2006) demon-
strated that Ara h 6 is a heat and digestive stable protein resis-
tant to proteolytic treatment. Ara h 7 also belongs to the
conglutin (2 S albumin) protein family and has a molecular
weight of 15 kDa, with 35% sequence homology to Ara h 2 and
Ara h 6 and recognized by 13% of the sera from 40 peanut-
allergic patients (Maleki et al., 2003). Currently, the WHO/
IUIS Allergen Nomenclature Subcommittee (2017) recognizes
two Ara h 7 isoforms, Ara h 7.0101 and Ara h 7.0201 (Table 1).
Other proteins (profilin, pathogenesis-related proteins,
nonspecific lipid transfer proteins, oleosin, defensins)
Ara h 5 with a molecular weight of 15 kDa belongs to the profi-
lin family of proteins (Table 1) and functions to regulate the
polymerization of actin (Breiteneder and Radauer, 2004;
WHO/IUIS Allergen Nomenclature Sub-Committee, 2017).
Ara h 5 is present in low levels in peanut extracts and is only
IgE reactive with approximately 13% of 40 peanut-allergic
patients (Zhou et al., 2013).
Ara h 8 with a molecular weight of 17 kDa has been identified
as a pathogenesis-related protein (Table 1) produced in plants in
the event of pathogen invasion and serve as host protection (Van
Loon, 1985). Ara h 9 functions as a non-specificlipid-transferpro-
tein to shuffle phospholipids and other fatty acid groups between
cell membranes with a molecular weight of 9.8 kDa (S
aiz et al.,
2013). However, few allergen studies have been conducted to bio-
chemically characterize the allergenicity of Ara h 9.
Ara h 10 (16 kDa) and Ara h 11 (14 kDa) belong to the
oleosin structural protein family (Table 1) and are found in the
plant oil bodies and serve to stabilize the oil body structure dur-
ing maturation (S
aiz et al., 2013). Due to their association with
oil bodies, Ara h 10 and Ara h 11 extraction and isolation are
complicated and proven unsatisfactory (Millichip et al., 1996).
Ara h 12 and Ara h 13 proteins are peanut defensins (Table 1)
with molecular weights ranging from 5 to 12 kDa (WHO/IUIS
Allergen Nomenclature Subcommittee, 2017), which function
as host defense peptides against bacteria and/or fungi (Bublin
and Breiteneder, 2014).
Lipids
Most peanuts grown worldwide are primarily produced for edi-
ble oil, due to its desirous mild taste and high smoke point rela-
tive to other vegetable-based cooking oils. After oil extraction
from the seed, the remaining peanut meal is approximately
50% protein (Zhao et al., 2012).
Highly processed peanut oil (acid extracted, heat distilled)
has been shown not to contain peanut proteins and can be
safely consumed by peanut allergic subjects. However, cold-
pressed or cold-extruded peanut oils, processed at lower tem-
peratures may contain traces of peanut protein and may elicit
allergic responses in peanut-sensitive patients (du Plessis and
Steinman, 2004). Aflatoxin, a carcinogenic compound pro-
duced by the two fungi, Aspergillus flavus and Aspergillus para-
siticus, are generally not found in refined peanut oil.
However, aflatoxin contaminants can be found in crude or
lightly processed peanut oil (Sanders, 2002).
In many countries, peanuts seeds provide a significant nutri-
tious contribution to the diet due to their rich protein, lipid, and
fatty acid content. Traditional peanut seed ranges in oil content
from 44 to 56% with an average of 50% (Cobb and Johnson,
1973;Grossoetal.,1997), with a light yellow color and slightly
nutty flavor. Flavor and quality of peanuts and peanut products
are largely a function of the seed lipid chemistry, while peanut
lipid and fatty acid composition is greatly dependent upon culti-
var, seed maturity, and environmental conditions and geographic
location (Brown et al., 1975;Young,1996).
The chemical and physical properties of fats and oils are
mainly determined by the fatty acid profile of the oil and their
position within the triacylglycerol molecule. Peanut oil has a
high oleic content, which is associated with good oxidative and
frying stabilities. Peanut oil is a non-drying oil, which does not
harden when exposed to air and solidifies from 0 to 3C
(Young, 1996;O’Brien, 2004). The three major fatty acids pres-
ent in peanut oils as acylglycerols, esters formed from glycerol
and fatty acids are Palmitic (C16:0), Oleic (C18:1), and linoleic
(C18:2) acids. Normally Steric (C18:0), Arachidic (C20:0), Eico-
senoic (C20:1), Behenic (C22:0), and Lignoceric (C24:0) acids
occur in minor proportions, while trace levels of linolenic
(C18:3) can also be present (Carrin and Carelli, 2010).
CRITICAL REVIEWS IN FOOD SCIENCE AND NUTRITION 3
Peanut cultivars from the United States, Argentina, Bolivia,
and Poland have the following fatty acid distributions: C16:0 D
9.3 to 13.0%, C18:0 D1.1 to 3.6%, C18:1 D35.6 to 58.3%,
C18:2 D20.9 to 43.2%, C20:0 D0.3 to 2.4%, C20:1 D0.7 to
3.2%, C22:0 D1.8 to 4.4%, and C24:0 D0.4 to 1.9% (Branch
et al., 1990; Grosso and Guzman, 1991; Grosso et al., 1994). In
parallel, African peanut oils contain 44.5, 32.3, and 13.9% of
C18:1 (Oleic), C18:2 (Linoleic), and C16:0 (Palmitic), respec-
tively (Carrin and Carelli, 2010). Studies by Hinds (1995) dem-
onstrated that peanut seeds increased oleic fatty acid content,
while decreasing palmitic and linoleic fatty acids during
maturation.
Within the last decade, genetic manipulations have been
utilized to alter peanut chemistry and improve nutritional
quality of peanuts and peanut products. The incorporation of
high-oleic traits into peanut breeding lines has resulted in ele-
vated high-oleic fatty acid content and extended shelf life.
Normal-oleic conventional peanut cultivars have a lipid profile
of 52% oleic fatty acid and 27% linoleic fatty acids, while new
high-oleic peanut cultivars have a lipid profile of 80% oleic
fatty acids and 2% linoleic fatty acids (Isleib et al., 2006). The
primary difference found in high-oleic peanuts is the replace-
ment of linoleic acid by oleic fatty acids in the peanut oil.
Most of the fatty acids present in peanut oil are present as
triacylglycerols (TAG) at approximately 93.3 to 95.8% of weight
(Sanders, 2002). Studies by Sanders (2002) demonstrated that
TAG content is dependent upon seed maturation and increases
incrementally until full maturation. Sanders (2002) also con-
cluded that not only does environment and location affect fatty
acid composition, but also affects the spatial arrangement of
these fatty acids found within specific TAG molecules, with a
higher percentage of oleic or linoleic fatty acids in the sn-2 posi-
tion. Moreover, the spatial arrangement of these fatty acids
within the glycerol skeleton is nutritionally important. During
the digestive process, fatty acids found within the sn-2 position
are conserved, while fatty acids at the sn-1 and sn-3 positions
are released by pancreatic lipase (Carrin and Carelli, 2010).
Therefore, long-chain saturated fatty acids present preferen-
tially at these positions and with melting points higher than
human body temperature (C18:0, C20:0, C22:0, C24:0) remain
free and solid within the intestinal lumen with weak intestinal
absorption and therefore have no effect on plasma lipids
(Dubois et al., 2007).
Free fatty acids (FFA) and diacylglycerols (DAG) can also be
found in unprocessed peanut oil. Crude peanut oil can have an
FFA content as low as 0.3%, while commercial oil ranges from
0.5 to 1.5% (Padley et al., 1994). FFA and DAG levels within
peanut oil vary and are dependent upon seed maturity. For
example, the Florunner peanut variety has a reduction in FFA
content from 4.5 to 0.7%, and DAG levels from 2.4 to 0.5%,
between the white immature stage to full maturity (Ayres, 1983).
Healthy mature peanut seeds have an FFA content of less than
0.5% (Carrin and Carelli, 2010). However, if the seeds are dam-
aged (i.e., mold), FFA content up to 5% may be found (Ayres,
1983). Thus, high levels of FFA may indicate poor handling,
immaturity, or mold growth (Sanders et al., 1992).
Phospholipid content in peanut oil is very low (0.3 to 0.7%)
and is a major constituent of the cell membranes of the seed.
Peanut phospholipids (PL) have a high degree of unsaturation
and the major PL in conventional peanut oils are phosphatidyl-
choline, phosphatidic acid, phosphatidylethanolamine, phos-
phatidylinositol, and phosphatidylglycerol (Singleton and Sti-
keleather, 1995).
Carbohydrates
Oil or dry roasted peanuts contain approximately 21.51 g of
carbohydrates per 100 g (USDA, Food Composition Data-
base, 2017) with starch as the major carbohydrate. How-
ever, peanut research has demonstrated that peanut
carbohydrate content is dependent upon cultivar, matura-
tion, and geographic location (Pattee and Young, 1982)and
may contain the following carbohydrates in varying quanti-
ties (major to minor): sucrose, fructose, glucose, inositol,
raffinose, stachyose. Pattee and Young, (1982) reported that
upon thermal processing (roasting), sucrose undergoes
hydrolysis liberating fructose and glucose, which in turn
react with free amino acids to form the characteristic flavor
of roasted peanuts (Pattee and Young, 1982). Defatted pea-
nut flour has been shown to contain approximately 38%
total carbohydrates (Fig. 1) of which account for oligosac-
charides 18%, starch, 12.5%, hemicellulose A 0.5%, hemicel-
lulose B 3.5%, and cellulose (fiber) 4.5% (Tharanathan
et al., 1975). Of the oligosaccharide fraction, approximately
13.90% sucrose, 0.89% raffinose, 1.56% stachyose, and
0.41% verbascose in unprocessed peanut flour (Tharanathan
et al., 1975). Raffinose, stachyose, and verbascose, non-
digestible short-chain oligosaccharides (Fig. 1), are indigest-
ible in the human digestive tract and pass unchanged to the
colon, and are subject to bacterial fermentation in the lower
gut causing abdominal bloating (Li et al., 2013). The
enzyme alpha-galactosidase responsible for their digestion
and can be purchased as an over-the-counter supplement to
prevent gas after eating legumes.
Starch is a homopolysaccharide made up of a-D glucose res-
idues joined by glycosidic bonds. Upon digestion, salivary and
pancreatic amylase catalyzes the hydrolysis of starch to maltose
and maltotriose, isomaltose (Zeeman et al., 2010). Subse-
quently, these disaccharides are catalyzed by digestive enzyme
sucrose-isomaltase (Fig. e 1) found within the apical brush bor-
der membrane of the small intestine to liberate two units of glu-
cose (Gericke et al., 2017), the functional unit of energy needed
to fuel growth, development, and maintenance.
Pattee et al. (2000) demonstrated that carbohydrate compo-
sition is also dependent upon market-type and maturation.
However, in these studies while the carbohydrate values were
different between market-types, they were not significantly dif-
ferent. Moreover, studies by Pattee et al. (1998) reported that
increased sweetness directly related to carbohydrate content
was associated with superior flavor profiles, with reduced bit-
terness and improved roasted peanut flavor.
Micronutrients
Vitamins
Peanuts provide a valuable source of water-soluble B vitamins
and Vitamin E (tocopherol). Vitamin E (tocopherol) is a fat-
4O. T. TOOMER
soluble vitamin that is an antioxidant. Tocols (fundamental unit
of the tocopherol family) are naturally occurring antioxidants
found in plant oils like peanuts and include four tocopherol and
four tocotrienol (members of the Vitamin E family) isomers, des-
ignated as a,b,g,andd. These antioxidants inhibit lipid peroxi-
dation in foods by stabilizing hydro-peroxides and other free
radicals. Studies have demonstrated that these antioxidants
decrease during processing of the peanut oil, with chemical refin-
ing removing as much as 10 to 20% of the tocopherols and toco-
trienols, and 30 to 60% being lost with deodorization or steam
distillation (O’Brien, 2004). Crude peanut oil has only 30 to 40%
of the tocopherol content that soybean oil has, but has almost
three times as much tocotrienols than soybean oil (Carrin and
Carelli, 2010). Studies by Chun (2002) reported a tocopherol
content of 8.2 mg/ 100 g in raw conventional peanuts and a
tocopherol content of 4.1 mg/100 g in roasted peanuts (Table 2).
Hashim et al. (1993) found that tocopherol content in pea-
nut oil was dependent upon stage of maturation and peanut
cultivar, with there being significant differences between matu-
rity stages of runner- and Virginia-type peanut cultivars. Upon
comparisons of tocopherol content in peanut oil from peanuts
grown in Argentina, China, and the United States, Sanders
et al. (1992) found that peanut oil from peanuts grown within
the United States had higher contents of tocopherols (210.1 to
243.8 ppm) and lowest in peanut oil from peanuts grown in
China (102.9 to 183.9 ppm). Therefore, peanuts can provide a
dietary source of vitamin E important to human nutrition.
Additionally, peanuts are a good source of water-soluble
vitamin thiamine (B
1
), which functions as a coenzyme in carbo-
hydrate and amino acid metabolic pathways (Table 2). Studies
by Dougherty and Cobb (1970a) reported thiamine content in
the peanut seed to be about 1.0 mg/100 g, while thiamine con-
tent in peanut testa (skin) to be considerably higher at approxi-
mately 3.8 mg/100 g (Dougherty and Cobb, 1970b). Peanuts
are also an efficient source of riboflavin (B
2
), which functions
as a coenzyme in carbohydrate, lipid, and protein metabolic
pathways and is approximately 0.098 mg riboflavin/100 g of
dry roasted peanuts (Settaluri et al., 2012). Peanuts also provide
B vitamin, niacin (B
3
) an essential coenzyme in metabolic
respiratory pathways within the mitochondria with an approxi-
mate amount of 13.525 mg/100 g of dry roasted peanuts
(Table 2, Settaluri et al., 2012).
Vitamin B
5
, pantothenic acid is responsible for the for-
mation of Coenzyme A, which is responsible for vital reac-
tions in energy metabolism, synthesis of cholesterol, and
synthesis of heme. Vitamin B
5
is present in peanuts at
approximate amounts of 1.395 mg/100 g of dry roasted pea-
nuts (Table 2, Settaluri et al., 2012). Additionally, in smaller
amounts vitamin B
6
(pyridoxine) and vitamin B
9
(folic
acid) can be found in peanuts with approximate amounts
of 0.256 mg and 145 mgper100gofdryroastedpeanuts,
respectively (Table 2, Settaluri et al., 2012). Vitamin B
6
(pyridoxine) functions biologically as an essential coenzyme
in amino acid, glucose, and lipid metabolism pathways.
Moreover, vitamin B
6
(pyridoxine) is needed for neuro-
transmitter, histamine, and hemoglobin synthesis, while
vitamin B
9
(folic acid) is an essential vitamin needed for
RNA, DNA synthesis, amino acid metabolism, cell division,
and fetal development.
Minerals
Peanuts are a good dietary source of the macro minerals
(Derise et al., 1974; Settaluri et al., 2012), which are the miner-
als needed daily in a quantity greater than 100 mg/day. Studies
by Derise et al. (1974) demonstrated only slight variations in
the mineral content between various peanut cultivars (Virgina-
70, NC-2, Florigiant) and between raw and roasted peanuts.
Peanuts contain approximately 658 mg/ 100 g (Table 2, Setta-
luri et al., 2012) of the vital mineral potassium, which functions
along with sodium to maintain the bodies’electrolyte balance,
and muscle and neurological function. Settaluri et al. (2012)
also demonstrated that peanuts provide the macro minerals
magnesium (175 mg), calcium (54 mg), and phosphorus (358 mg)
per 100 g of dry roasted peanuts (Table 2). Magnesium is needed
for normal muscle and nerve function and maintenance of blood
pressure, while calcium is required for normal bone and tooth
development and muscle function. Phosphorus (358 mg) along
Figure 1. Carbohydrates in peanut flour from whole conventional peanuts. Defatted contains approximately 38% total carbohydrates comprising 18% oligosaccharides,
12.5% starch, 0.5% hemicellulose A, 3.5% hemicellulose B, and 4.5% cellulose (fiber) (Tharanathan et al., 1975). The oligosaccharide fraction is comprised of 13.90%
sucrose, 0.89% raffinose, 1.56% stachyose, and 0.41% verbascose. Raffinose, stachyose, and verbascose, non-digestible oligosaccharides are indigestible in the human
digestive tract and are subject to bacterial fermentation in the lower gut causing abdominal bloating. Starch is a homopolysaccharide made up of a-D glucose residues
joined by glycosidic bonds. Salivary and pancreatic amylase catalyzes the hydrolysis of starch to maltose and maltotriose. Subsequently, these disaccharides are catalyzed
by digestive enzyme maltase, sucrose, and/or sucrose-isomaltase to free glucose.
CRITICAL REVIEWS IN FOOD SCIENCE AND NUTRITION 5
with calcium is required for bone and teeth formation, and protein
synthesis in tissue growth and repair. Peanuts contain approxi-
mately 5.56 mg of sodium per 100 g of roasted peanuts (Table 2.
Derise et al., 1974). Dietary sodium is important biologically for
electrolyte balance, hydration, and proper functioning of nerves
and muscles. In general, peanuts are a good dietary source of
potassium, phosphorus, and magnesium.
Peanuts also provide a source of trace minerals, which are
minerals needed daily in a quantity less than 100 mg/day. Setta-
luri et al. (2012) reported that peanuts provide the trace
Table 2. Micronutrients content of whole conventional peanuts.
Vitamins Class Name Peanut content Main biological function References
Fat soluble Tocopherol 8.2 mg/ 100 g raw, 4.1
mg/ 100 g roasted
Antioxidant Chun, 2002
Water soluble Thiamine (B1), 1.0 mg/100 g peanut seed coenzyme in carbohydrate and
amino acid metabolic
pathways
Dougherty and Cobb,
1970a
Riboflavin (B2) 0.098 mg/100 g of dry roasted
peanuts
coenzyme in carbohydrate, lipid,
and protein metabolic
pathways
Settaluri et al., 2012
Niacin (B3) 13.525 mg/100 g of dry roasted
peanuts
coenzyme in metabolic
respiratory pathways
Settaluri et al., 2012
Pantothenic acid
(B5)
1.395 mg/100 g of dry roasted
peanuts
formation of Coenzyme A Settaluri et al., 2012
Pyridoxine (B6) 0.256 mg/100 g of dry roasted
peanuts
coenzyme in amino acid,
glucose, and lipid
metabolism pathways
Settaluri et al., 2012
Folic acid (B9) 145 mg/100 g of dry roasted
peanuts
nucleic acid synthesis, amino
acid metabolism, early
development
Settaluri et al., 2012
Minerals Macro minerals Potassium 658 mg/ 100 g of dry roasted
peanuts
electrolyte balance, and muscle
and neurological function
Settaluri et al., 2012
Magnesium 175 mg/ 100 g of dry roasted
peanuts
muscle and nerve function and
maintenance of blood
pressure
Settaluri et al., 2012
Sodium 5.56 mg/ 100 g of roasted
peanuts
electrolyte balance, hydration,
function nerves and muscles
Derise et al., 1974
Calcium 54 mg/ 100 g of dry roasted
peanuts
normal bone and tooth
development and muscle
function
Settaluri et al., 2012
Phosphorus 358 mg/ 100 g of dry roasted
peanuts
bone and teeth formation, tissue
growth and repair
Settaluri et al., 2012
Trace minerals Zinc 3.31 mg/100 g of dry roasted
peanuts
immune system function,
wound healing, cell division,
and growth
Settaluri et al., 2012
Iron 2.26 mg/ 100 g of dry roasted
peanuts
essential element for blood
production and oxygen
transfer
Settaluri et al., 2012
Manganese 2.06 mg/ 100 g of roasted
peanuts
formation connective tissues,
blood clotting factors, sex
hormones, nerve function
Derise et al., 1974
Copper 0.671 mg/ 100 g of dry roasted
peanuts
red blood cells, healthy blood
vessels, nerves, bones,
immune support
Settaluri et al., 2012
Selenium 7.5 mg/ 100 g of dry roasted
peanuts
antioxidant, prevent cell
damage
Settaluri et al., 2012
Table 3. Phytonutrient content of whole conventional peanuts.
Class Name Peanut content Biological function References
Isoflavonoid Daidzein 49.7 mg/ 100 g of dry roasted peanuts precursor S-equol a non-steroidal
selective agonist of the beta
estrogen receptor
Mazur et al., 1998;
Mazur, 1998
Genistein 82.6 mg/ 100 g of dry roasted peanuts antioxidant, anthelmintic, angiogenesis
inhibitor, inhibits cancer cell growth
Mazur et al., 1998;
Mazur, 1998
Phenolic acids p-coumaric acid 6.9 mg/100 g of dry roasted peanuts Talcott et al., 2005a,
2005b
Phytosterols b-sitosterol 61 mg to 114 mg/100 g of roasted
peanuts
may inhibit cancer growth, protect
against heart disease
Awad et al., 2000
Stilbenes Resveratrol 0.48 mg/g to 3.96 mg/g
*
peanut
content with abiotic stress
anti-obesity, anti-diabetic,
neuroprotective, cardioprotective,
chemo-protective
Rudolf and
Resurreccion, 2006
peanut content with abiotic stress
6O. T. TOOMER
minerals zinc (3.31 mg), iron (2.26 mg), copper (0.671 mg), and
selenium (7.5 mg) per 100 g of dry roasted peanuts (Table 2).
Derise et al. (1974) reported that peanuts provide approxi-
mately 2.06 mg of manganese per 100 g of roasted peanuts.
Zinc is important biologically for proper function of the
immune system, wound healing, and repair. Iron is an essential
element needed for heme blood production and the transfer of
oxygen within the body, while copper is needed for the forma-
tion of red blood cells and healthy blood vessels, nerves, and
bones. Manganese is biologically important for the formation
of connective tissue, bones, blood clotting factors, sex hor-
mones, and nerve function, while the trace mineral selenium
functions as an antioxidant. Based on these results, peanuts can
provide over 70% of the daily need for copper and 14% of the
daily need for selenium (Settaluri et al., 2012).
Phytonutrients
Isoflavonoids
Unlike macronutrients (proteins, fats, and carbohydrates), vita-
mins and minerals, phytonutrients are not essential to maintain
life. However, when consumed they may help in disease pre-
vention and promote health and wellness. Studies have demon-
strated that peanuts provide valuable sources of these
phytonutrients and therefore of nutritional importance as a
functional food (Mazur et al., 1998, Mazur 1998). Studies by
Mazur et al. (1998) and Mazur (1998) reported that peanut
seeds had an isoflavonoid content of daidzein and genistein in
the greatest amounts with a content of 49.7 mg/100 g and 82.6
mg/100 g, respectively (Table 3). Daidzein and genistein are iso-
flavone compounds that are commonly found in a number of
plants, and are particularly abundant in soybeans and soy prod-
ucts (USDA Database, 2008).
Genistein and daidzein have similar structures to human estro-
gen and therefore are both identified as phytoestrogens. In plants,
genistin is the precursor glucoside and in vivo is hyrdrolyzed by
digestive enzymes to genistein and glucose. Various studies have
demonstrated that genistein functions biologically also as an anti-
oxidant(Hanetal.,2009,Zhaoetal.,2016), anthelmintic (Tandon
et al., 2003), angiogenesis inhibitor (Farina et al., 2006), and inhib-
its cancer cell growth (Gossner et al., 2007;Raynaletal.,2008;
Nakamura et al., 2009;Kimetal.,2009). In vivo daidzein is metab-
olized by human intestinal bacteria to produce end metabolite, S-
equol, also known as (S)-(–)-4',7-isoflavandiol, an enantiomer of
the naturally occurring isoflavondiol estrogen a non-steroidal
selective agonist of the beta estrogen receptor (Muthyala et al.,
2004). Research studies have reported the use of S-equol for the
successful treatment of menopausal symptoms in women (Jackson
et al., 2011).
Phenolic acids
Phenolic acids are abundant in plant-based foods (seeds, skins
of fruits, leaves of vegetables) and therefore readily available in
a balanced diet. Upon consumption phenolic acids are readily
absorbed through the intestinal walls and may directly function
as an antioxidant by preventing cellular damage due to oxida-
tive free radicals (Gon¸calves et al., 2017; Viapiana and
Wesolowski, 2017). Moreover, studies have demonstrated that
dietary phenolic acids promoted anti-inflammation in LPS
challenged mice (Choi et al., 2017), in a murine model of
induced-colitis (Monk et al., 2016), and a murine model of
high dietary fat (Chang et al., 2015).
Normal oleic acid peanuts (raw and/or dry roasted) had
the highest content of polyphenolic compounds in compari-
son to mid and high oleic peanut varieties, with free p-cou-
maricacid,threeesterified derivatives of p-coumaric, and
two esterified derivatives of hydrobenzoic acid identified as
the predominate polyphenolic compounds found in peanuts
(Talcott et al., 2005a,2005b). Benzoic acid is a polyphenolic
compound with gallic acid and cinnamic acid as derivatives
(Natella et al., 1999), and p-coumaric as the hydroxyl deriva-
tive of cinnamic acid (Fig. 2). Whole raw peanuts had a
range of 8 mg/kg to 66 mg/kg if p-coumaric acid (Table 3)
among peanut cultivars, with the value increasing to an aver-
age of 69 mg/kg upon dry roasting (Table 3, Talcott et al.,
2005a,2005b). These studies demonstrate that differences in
the content of polyphenolic compounds between peanut cul-
tivars were highly dependent upon background genetics,
growth conditions, disease resistance, post-harvest handling,
and thermal treatment.
Phytosterols
Phytosterols, (i.e., plant sterols) are essential components of cell
membranes and are similar to cholesterol. b-sitosterol has been
shown to be the predominate source of phytosterol (PS) in pea-
nuts with a content of 61 mg to 114 mg/100 g of roasted pea-
nuts (Table 3; Awad et al., 2000). Studies have reported that
b-sitosterol consumption may inhibit cancer growth (Awad
et al., 1998; Roussi et al., 2005; Imanaka et al., 2008) and protect
against heart disease (Peanut Institute, 2000; Bouchenak and
Lamri-Senhadji, 2013). Among the peanut cultivars, Valencia
peanuts (raw, dry, or oil roasted) contained the highest phytos-
terol content, with peanut butter (144–157 mg/100 g) and pea-
nut flour (55–60 mg/100 g) containing significant amounts
of phytosterols as well (Awad et al., 2000). Unrefined peanut
oil has a phytosterol content of approximately 207 mg
Figure 2. Plant phenylpropanoid pathway-biosynthesis of p-coumaric acid. L-Phe-
nyalanine is an essential amino acids for mammals and dietary intake is necessary
for protein synthesis. Fungi and plants produce phenylalanine via the shikimic acid
pathway. Phenylalanine is directly utilized for protein synthesis in plants or metab-
olized through the phenylpropanoid pathway. Phenylalanine ammonia-lyase (PAL)
is the enzyme responsible for the conversion of phenylalanine to intermediate cin-
namic acid. Cinnamic acid-4-hydroxylase (CA4H) catalyzes the conversion of cin-
namic acid to p-coumaric.
CRITICAL REVIEWS IN FOOD SCIENCE AND NUTRITION 7
phytosterols/100 g, which is 38% higher than the phytosterols
content found in olive oil (Awad et al., 2000). Studies by Koeh-
ler and Song (2002) also determined that phytosterol content
was not only influenced by cultivar, but also by maturation,
with increasing amounts with maturity.
While phytosterols are similar to cholesterol, they are poorly
absorbed across the intestinal mucosa and therefore circulating
concentrations are low. After uptake by the intestinal entero-
cytes, phytosterols are actively excreted back into the intestinal
lumen by the cassette (ABCG5/G8) transporter (Nissinen et al.,
2002). It has been documented that phytosterols displace die-
tary cholesterol from uptake during the digestive process and
therefore reduce the uptake and levels of circulating cholesterol
(Nissinen et al., 2002). In a placebo-controlled study, moderate
(0.46 g/day) and high (2.1 g/day) consumption of phytosterols
reduced cholesterol absorption by about 10 and 25%, respec-
tively, while significantly increasing excretion of endogenous
and dietary cholesterol by 36 and 74%, respectively (Racette
et al., 2010).
Stilbenes
Stilbenes chemically contain two phenyl compounds joined by
a two-carbon methylene bridge. Stilbenes are a member of the
vast group of polyphenols naturally occurring in plants. Stil-
benes, like isoflavonoids are classified as phytoestrogens. Res-
veratrol (3,5,40-trihydroxy-trans-stilbene), the most extensively
studied stilbene, is a polyphenol phytoalexin mainly found in
the skin of grapes. Recently, resveratrol has attracted tremen-
dous scientific interest due to its potential health benefits
related to cardiovascular (Diaz et al., 2016), chemo-protective
(Zhang et al., 2015), anti-obesity (Zou et al., 2017), anti-diabetic
(Tran et al., 2017), and neuroprotective (Pineda-Ram
ırez et al.,
2017) properties.
Stilbenes found in peanut include resveratrol, 3-isopenta-
dienyl resveratrol, and various arachidins (Ku et al., 2005). Res-
veratrol in peanuts serves to protect the plant from plant
pathogens (Higgs, 2003). While peanuts are a source of resvera-
trol, the content is low in comparison to that of grapes. How-
ever, resveratrol content is much greater in the leaves, roots,
and shells of peanuts (Chung et al., 2003). Seo et al. (2005)
reported that resveratrol peanut content was dependent upon
post-harvest processing, with a 45- to 65-fold increase after a
treatment of 20-hour soaking/66-hours drying. Other studies
by Rudolf and Resurreccion (2006) reported significant
increases (0.48 mg/g to 3.96 mg/g) in resveratrol content with
abiotic stresses (Table 3). Boiled peanuts having 10-fold greater
resveratrol content in comparison to roasted peanuts or peanut
butter (Rudolf, 2003).
Upon consumption, dietary resveratrol is well absorbed
across the intestinal mucosa; however, its bioavailability is low
due to its rapid metabolism and elimination. Resveratrol is
rapidly metabolized by conjugation to glucuronic acid and/or
sulfate, forming resveratrol glucuronides, sulfates, and/or sul-
foglucuronides, with these sulfate conjugates being the major
metabolites found in the plasma and urine of humans
(Burkon and Somoza, 2008). Administration of single oral
doses of 25 mg of trans-resveratrol to healthy volunteers
resulted in peak blood concentrations of total resveratrol (i.e.,
trans-resveratrol plus its metabolites) around one hour later,
at approximately 1.8–2mmoles/l (mM), depending on whether
resveratrol was administered in wine, vegetable juice, or grape
juice (Goldberg et al., 2003; Walle et al., 2004).
Conclusion
Prior to the United States Civil War, peanuts were considered a
regional Southern food. After the United States Civil War, tech-
nological advancements led to increased demand for peanut oil,
peanut butter, and peanut products (roasted and salted and con-
fections). Additionally, the scientific discoveries of George Wash-
ington Carver identified numerous non-food uses of the peanut
and peanut plant, which encouraged the cultivation of peanuts
as a profitable rotational crop for cotton (Agricultural Marketing
Resource Center, 2017). Peanuts and peanut butter became an
integral part of the American Armed Forces rations during
World War I and II, with the United States Army popularizing
the peanut butter and jelly sandwich for sustenance during mili-
tary maneuvers in World War II (National Peanut Board, 2017).
Today, peanuts are the 12th most valuable cash crop grown
in the United States with an estimated farm value over one bil-
lion U.S. dollars (American Peanut Council, 2017). Americans
eat more than six pounds of peanut products annually, valued
at more than $2 billion retail, with peanut butter accounting
for approximately $850 million (American Peanut Council,
2017). Peanut butter has become an extremely popular, nutri-
tious, and economical sandwich spread for children and adults
across the United States.
As with many other food items, interest in nutritional com-
position and chemistry is resultant of their use in human food.
Over the years, peanut feeding studies have demonstrated that
regular peanut consumption has been linked to reduced heart
disease (Jones et al., 2014), certain types of cancers (Gonzalez
and Salas-Salvad
o, 2006), and improved weight management
(Moreno et al., 2013). Moreover, other studies have identified
the value in the phytonutrient composition of peanuts, which
may improve overall health and wellness (Bouchenak and
Lamri-Senhadji, 2013; Sales and Resurreccion, 2014).
An improved knowledge or better understanding of the
nutritional chemistry of peanuts enables us to better harness
the power of these nutrients in improved peanut products
within the food industry. Moreover, improved understanding
of the nutritional chemistry of peanuts may help to identify the
use of peanuts and/or peanut components more effectively in
the agricultural feed industry to improve the health, growth,
and performance of production animals.
While improved comprehensive understanding of the nutri-
tional chemistry of peanuts better enables us to not only
address issues related to nutrition and hunger worldwide, but
also potentially improve health and wellness of consumers by
knowledge of the functional components found within peanuts.
Therefore, the spectrum of new and emerging peanut nutri-
tion-related research continues to greatly expand. This review
provides substantial evidence that peanuts are a nutritious food
and food ingredient packed with health-promoting bioactive
compounds and worthy of additional nutrition-related experi-
mentation regarding this valuable agricultural commodity.
8O. T. TOOMER
Acknowledgments
The author would like to thank Sabrina Whitley-Ferrell, in the Market
Quality and Handling Research Unit for her assistance with the prepara-
tion of this manuscript. The author has no conflicts of interest to declare.
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