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Czech J. Food Sci., 35, 2017 (2): 95–105 Review
doi: 10.17221/114/2016-CJFS
White Lupin (Lupinus albus L.) – Nutritional and Health
Values in Human Nutrition – a Review
Janusz Prusinski*
Department of Agrotechnology, UTP University of Science and Technology, Bydgoszcz, Poland
*Corresponding author: janusz.prusinski@utp.edu.pl
Abstract
Prusinski J. (2017): White lupin (Lupinus albus L.) – nutritional and health values in human nutrition – a review.
Czech J. Food Sci., 35: 95–105.
White lupin seeds have been used in human nutrition and treatment for several thousand years. Nowadays the use of
white lupin seeds is limited by a small scale of their production. However, in the last 20 years quite new properties of
white lupin have been discovered for the application in the production of different kinds of functional food. Unique
traits of protein, fatty acids with a desirable ratio of omega-6 to omega-3 acids, and fibre as well as other specific
components, for example oligosaccharides and antioxidants or non-starch carbohydrates, make white lupin an excel-
lent component in many healthy diets. The effects of white lupin components concern the physiological condition of
the human body, including diabetes, hypertension, obesity, cardiovascular diseases, lipid concentration, glycaemia,
appetite, insulin resistance, and colorectal cancer. Seeds are used among others for the production of gluten-free flour,
bacterial and fungal fermented products, noodle and pasta products, as substitutes of meat, egg protein and sausages,
also are cooked, roasted and ground and mixed with cereal flour in the production of bread, crisps and pasta, crisps
and dietary dishes.
Keywords: health benefits; chemical composition; effects on human health
Many researchers state that the use of lupin seeds
for consumption and medicinal purposes has been
the subject of interest for more than 3000 years
around the Mediterranean Sea. White lupin is the
longest known crop species in the history of the
genus Lupinus – it was known among Aegean farm-
ers at least 400 years B.C. Until the beginning of the
19th century in Europe – in the Mediterranean Sea
region it was the most often cultivated lupin species
for green manure and for seeds, which were used in
animal and human nutrition. However, because of
high alkaloid content, the seeds could not be con-
sidered as a safe food component (Prusinski 2015).
A traditional way to avoid bitter taste caused by the
presence of alkaloids in the lupin seeds was their very
fine grinding, and next multiple rinsing with water,
which however caused a decrease in the nutritive
value as a result of removing soluble proteins, free
amino acids, carbohydrates, and minerals. Neverthe-
less, this method was commonly used by societies
in Mediterranean Europe and Andean countries
(Petterson 1998).
After a significant success in qualitative cultiva-
tion in the 20th century (Prusinski 2015) a signifi-
cantly increased interest was observed concerning
white lupin among feed and food producers, and
also, among others, in medicine, food processing,
and even in cosmetic industry, in the production of
ecological pesticides, etc. Laboratory tests were very
promising, however, the commercial use of white
lupin seeds for the above-mentioned purposes is still
insufficient and limited (Swan 2000). Meanwhile,
due to the development of lifestyle diseases caused
by an improper diet, i.e. cardiovascular diseases,
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obesity, and diabetes, special attention is paid to
healthy nutrition. White lupin seeds provide numer-
ous health benefits counteracting the problems which
are also connected with high blood pressure, insulin
resistance, or higher cholesterol level. Taking into
consideration predictions concerning a continuously
growing number of deaths as a result of lifestyle
diseases, functional food, including the use of lupin
seeds, will have a greater and greater importance
(Martins & Bento 2007; Martirosyan & Singh
2015). In many countries of Europe, the conservative
attitude of consumers towards genetically modified
organisms inhibited work on the popularisation
of GMO in a significant way, and caused aversion
towards importing products obtained from GMO
cultivars (Sweetingham & Kingwell 2008). In
the future, the lack of GMO lupin may lead to a sig-
nificant development of a functionally varied food,
mainly from white lupin seeds.
Basic seed composition and properties
of its constituents
White lupin may be cultivated on almost all conti-
nents, however, chemical composition of seeds and
their nutritive value depend on plant response to
environmental conditions. Only the protein content
does not significantly depend on the place of cultiva-
tion, while the proportion of other components in
seeds is significantly dependent on the cultivation
region (Bhardwaj et al. 1998).
Protein. Owing to symbiosis with rhizobia, white
lupin has an ability to fix N2
with the use of atmo-
spheric nitrogen for the production of protein and
other nitrogenous substances in seeds, which contain
hardly any starch at the same time (Kurlovich et
al. 2002).
The average protein content in the seeds of white
lupin is from 32.9% (Martinez-Villaluenga et
al. 2006; Strakova et al. 2006) up to more than
36.0% (Sujak et al. 2006), or even 38.0% (Vecerek
et al. 2008; Saastomoinen et al. 2013). Storage
proteins in lupin consist of 85% globulins and 15%
albumins (Petterson 1998). The globulin frac-
tion contains 3 main proteins of varied amino acid
composition: α-, ß-, and γ-conglutin. γ-Conglutin
containing more methionine, cysteine, and valine
is a sulphur-containing amino acid that constitutes
about 4% of the protein composition (Duranti et
al. 1981). Some of them may have an allergenic effect
(Guillamon et al. 2010), however, compared to other
legumes (peas, soybean, bean), white lupin seeds have
a minimum content of proteins with anti-nutritive
properties (Kurlovich et al. 2002) and a higher
content of arginine, lysine, leucine, and phenylalanine
than for example soybean, which makes white lupin
seeds more valuable than other species regarding
nutrition standards (Table 1). White lupin contains
more amino acids (AA), including essential amino
acids (EAA), and is also characterised by a higher
index of essential amino acids (EAAI) and protein
efficiency ratio (PER), as well as a higher nutritive
value of protein isolates (CS) than the other two
lupin species (yellow and narrow-leafed) cultivated
in Europe (Sujak et al. 2006). Such composition
indicates a high suitability of white lupin protein for
vegans or for those on ovo-lacto vegetarian diet or
on gluten-free diet for people with the celiac disease
(Arnoldi & Greco 2011).
Dehulling of white lupin seeds increases crude
protein content and contents of most amino acids
(except alanine) (Straková et al. 2006) leading to
an increase of their biological value both for human
nutrition and for animal feeding. On the other hand,
extrusion of seeds leads to a decrease of amino acid
content and the value of EAA (Kiczorowska &
Lipiec 2002).
High-protein diet may play an important role in
ageing processes, pregnancy and lactation, growth
of youth and athletes, as well as in recovery. Some
white lupin proteins or their parts per se may posi-
tively affect the physiological condition of human
body under a wide range of unfavourable conditions,
including diabetes, hypertension, obesity, cardiovas-
cular diseases, or they may control glucose content
in diabetics or pre-diabetics (Duranti 2011).
Lupin flour contains approximately 30–40% protein,
and effectiveness of its utilization is slightly lower than
that of animal protein, though still satisfactory. The
human diet which is high in lupin proteins affects a
significant decrease in serum cholesterol, including
the LDL (low-density lipoprotein, bad cholesterol)
level, and also the level of triglycerides and glucose,
and it also lowers the blood pressure (Arnoldi 2005;
Nowicka et al. 2006). According to Naruszewicz
et al. (2006), thanks to white lupin addition to food,
a decrease in glucose content in the blood was 6.7%,
homocysteine 11.8%, and high-sensitivity C-reactive
protein (Hs-CRP) up to 18.3%.
The favourable effect of soybean protein is already
known, it can systematically decrease cholesterol
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content in humans and animals, and also reduce
the risk of the ischaemic heart disease. However,
modern industry is searching for products of specific
properties: technological and sensory ones, without
phytoestrogens and affordably priced. Clinical tests
proved capabilities of protein isolated from white
lupin to reduce cholesterol content to the same ex-
tent as it is observed in the case of other legumes
(Sirtori et al. 2004). The absence of phytoestrogens
in white lupin seeds makes this species better for the
young (Sirtori & Naruszewicz 2005).
Fat. Total fat content in white lupin seeds is from
about 8% (Straková et al. 2006; Uzun et al. 2007;
Vecerek et al. 2008; Andrzejewska et al. 2016)
to 11.5% (Sujak et al. 2006). In the subcontinen-
tal climate white lupin seeds are characterised by
an approximately 8% lower fat content than in the
Mediterranean climate (Annicchiarico et al. 2014).
White lupin seeds are an interesting source of
favourable ratios of important fatty acids used in
the prophylaxis of circulatory system diseases (Si-
mopoulos 2003). In their development, people
have consumed fats in which the ratio of ω-6 to
ω-3 acids was similar, whereas now in the diet of
highly-developed countries it is up to 15 to 1. Excess
ω-6 acids in the diet constitute a risk factor, while
the 2 : 1 ratio, which is observed in white lupin seeds,
has an enormous effect and decreases mortality as-
sociated with circulatory system diseases. According
to Green and Oram (1983), Arnoldi and Greco
(2011), and Andrzejewska et al. (2016) oleic acid
(mostly > 50%) prevails in the fat of white lupin
seeds, which indicates a high suitability of lupin oil
for consumption purposes (Table 2). In the seeds
of bitter white lupin cultivars (spp. termis) the pro-
portion of oleic acid is significantly higher (52.2 ±
Table 1. Amino acid composition of white lupin seeds (g/16 g N)
Specification
Sujak et al. (2006)
(Europe)
Petterson & Fairbrother (1996)
(Australia)
Bhardwaj (2002)
(USA)
Essential amino acids
Lysine 4.9 4.31 1.19
Methionine + cysteine 2.5 1.96
Cysteine 1.9 0.63
Threonine 3.5 3.16 0.83
Isoleucine 4.3 3.69 1.26
Tryptophan 0.6 0.97
Valine 4.1 3.55 1.06
Leucine 7.8 6.05 1.92
Histidine 3.3 1.74 0.51
Phenylalanine 5.6 3.33 1.02
Phenylalanine + tyrosine 5.6 8.68
Tyrosine 1.7 1.32
Non-essential amino acids
Arginine 11.4 2.98
Asparagine 10.5 2.96
Serine 4.5 0.99
Glutamine 23.5 6.08
Proline 3.5 1.31
Glycine 4.3 0.92
Alanine 3.2 0.83
Nutritional values
Amino acids participation (g/16 g N) 97.7
Essential amino acid participation (g/16 g N) 33.5
Chemical score of restrictive amino acid(s) 73.8
Essential amino acids index 85.0
Protein efficiency ratio 2.87
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2.32%) than in sweet cultivars (spp. albus) (44.9 ±
2.16%) (Alamri 2012). Andrzejewska et al. (2016)
identified in white lupin seeds also the following ac-
ids: myristic, pentadecanoic, palmitoleic, margaric,
arachidic, eicosadienoic, eicosatrienoic, docosadi-
enoic, and lignoceric at a content not higher than
1% each. Fatty acid forms (S) in Turkey (Erbas et al.
2005) were similar to those obtained by Bhardwaj
(2002) in the USA.
Carbohydrates. Lupin seeds and seed coats con-
tain various types of carbohydrates, mainly non-
starch carbohydrates (Khan et al. 2015), which are
the most abundant in seeds. In the seed coat, apart
from a small amount of proteins and fats, structural
polysaccharides prevail: cellulose, hemicellulose, and
pectins, while in the cotyledons – non-structural
polysaccharides of cell walls (Straková et al. 2006;
Vecerek et al. 2008), with the major proportion of
galactose, arabinose, and uronic acid (Mohamed
& Rayas-Duarte 1995; Petterson 1998). Most
carbohydrates represent soluble or insoluble fibre
up to about 2.83 g/100 g dry matter (Martinez-
Villaluenga et al. 2006).
Lupin starch is slowly digested and thus gradually
releases glucose into the blood. However, according
to some authors, mature air-dried white lupin seeds
do not contain any starch (Petterson 1998; Borek
et al. 2011) or its content is very low (Mohamed &
Ryas Duarte 1995; Martinez-Villaluenga et al.
2006) (Table 3). As a result, lupin seeds indicate a
low glycaemic index (Gullion & Champ 2002) and
can prevent diseases related to insulin resistance. In
Czech studies starch content in white lupin seeds
was at least twice higher (Straková et al. 2006;
Vecerek et al. 2008), which is however still very
low compared to starch content e.g. in pea seeds
(53.6–57.2%) (Dostálová et al. 2009).
Crude fibre. White lupin seeds are a valuable
source of dietary fibre (mostly insoluble), which is
higher than in soybean seeds (Pisarikova & Zraly
2010). In most Fabaceae plants the content of crude
fibre ranges from 8% to 27.5%, and that of soluble
fibre from 3.3% to 13.8% (Guillon & Champ 2002).
The average content of total fibre is from 101 g/kg
(Tizazu & Emire 2010) to 367 g/kg (Martinez-
Villaluenga et al. 2006) (Table 4).
The seed coat in white lupin, which constitutes
usually less than 20% of seed weight (Tizazu &
Emire 2010), is mainly composed of cellulose with
a small proportion of lignins. According to Pet-
Table 2. Triglyceride composition in the seeds of white lupin (%)
Fatty acids
Andrzejewska et al. (2015)
(Europe)
Alamri (2010)
(Asia)
Bhardwaj (2002)
(USA)
Oleic acid (18:1) (n-9) 54.3 44.9 50.9
Linoleic acid (18:2) (n-6) 14.9 26.2 23.5
Palmitic acid (16:0) 8.57 7.71 6.60
Linolenic acid (18:3) (n-3) 7.22 15.8 9.68
Gadoleic acid (C20:1) (n-9) 4.14
Erucic acid 1.59
Stearic acid (18:0) 1.57 1.71 0.78
Arachidic acids (20:0) and erucic acid (22:1) 0.81 + 1.59 2.74
Σ SFA (saturated fatty acids) 16.1 9.63 11.3
Σ MUFA (monosaturated fatty acids) 58.8 55.4
Σ PUFA (polyunsaturated fatty acids) 15.0 9.38 33.3
Total n-6 15.0
Total n-3 7.22
n-6/n-3 2.11
Table 3. Content of starch in white lupin seeds (g/kg)
Source Content
Mohamed & Ryas Duarte (1995) 30.0
Petterson (1998) not detected
Strakova et al. (2006) 81.5–85.7
Martinez-Villaluenga et al. (2006) 30.4
Vecerek et al. (2008) 72.6–86.5
Borek et al. (2011) not detected
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terson (1998), there is up to 95 g of fibre in 100 g
of the seed coat of white lupin. Thus, the dehulling
of white lupin seeds decreases the total fibre con-
tent and increases the soluble dietary fibre content
(Pisarikova & Zraly 2010). Fibre contained in the
cotyledons constituting about 30–40% of the germ
weight consists of insoluble materials of the cell walls
of a structure similar to pectins, which are soluble
fibre known for its favourable effect on lowering the
total cholesterol level as well as reducing susceptibil-
ity to the ischaemic heart disease (Hall et al. 2005).
Ash. White lupin seeds are a rich source of macro
and microelements (Table 5); their total content is
30–40 mg/kg (Straková et al. 2006; Sujak et al.
2006; Saastamoinen et al. 2013). Among macroele-
ments K, Mn, and Mg definitely prevail and among
microelements Ca, Fe, and Na are dominant (Ka &
Chandravanshi 2014). Lower Ca content improves
culinary properties of seeds and reduces the time of
their overcooking (Tizazu & Emire 2010).
Compounds of favourable/unfavourable effect
on human health
Anti-nutritional compounds. The negative effect
of unfavourable compounds in white lupin seeds
which is often cited in literature is outdated to a
significant extent now. As distinguished from other
legumes (peas, soybean, and bean), white lupin seeds
are characterised by a low or very low content of anti-
nutritive substances (Muzquiz et al. 1998; Enne-
king & Wink 2000). Their removal is possible either
through selection of genotypes with a low content of
these components or through post-harvest treatments,
e.g. germinating, cooking, soaking, fermentation, ex-
traction, etc.
Quinolizidine alkaloids (QAs) are a family of about
100 bitter components – secondary metabolites of
a bicyclic, tricyclic, and tetracyclic structure (Pet-
terson 1998). Seeds of wild lupin species may even
contain over 10 000 mg/kg alkaloids. The primary role
of alkaloids was to protect plants against herbivorous
animals. In their development, the alkaloid concen-
tration in different plant parts (leaves, roots, stems)
changes, reaching the highest value in the flowering
stage. In Australian studies on white lupin the oc-
currence of 4 alkaloids was observed: albine (15%),
13-hydroxylupanine (8%), lupanine (70%), and mul-
tiflorine (3%) in the total amount not reaching over
0.01% (Petterson & Fairbrother 1996), while in
Spanish studies additional a-isolupanine (Muzquiz
et al. 1994) as well as 13-α-angeloyloxylupanine and
13-tigloyloxylupanine were reported (Muzquiz et
al. 1998). Total alkaloid content in sweet white lu-
pin cultivars has been significantly reduced in the
process of domestication and breeding and does not
currently exceed 0.02% (Prusinski 2015).
Most alkaloids may cause cramps, vomiting, and
even death as a result of the respiratory system pa-
ralysis. Alkaloids also negatively affect the central
nervous system in mammals, though in very low
doses they may have a stimulating effect, while in
high doses an inhibiting effect (McKnickiene &
Asakaqviciute 2008). In humans too high dos-
age, especially of lupanine and sparteine, may also
cause trembling, arousal, and convulsions leading
to blurred vision, dry mouth, nervousness, and bad
mood (Arnoldi & Greco 2011). In many countries,
among others in France, Great Britain, Australia,
and New Zealand, the maximum alkaloid content
in flour and lupin products was established on the
level up to 200 mg/kg seeds (Resta et al. 2008). In
Table 4. Content of crude fibre in white lupin seeds (g/kg)
Source Content
Martinez-Villaluenga et al. (2006) 367
Sujak et al. (2006) 144
Straková et al. (2006) 107
Vecerek et al. (2008) 134
Tizazu & Emire (2010) 101
Table 5. Content of macro and minor elements of ash in
white lupin seeds (g/kg)
Element
Ka & Chandravanshi
(2014) (Africa)
Petterson & Fairbrother
(1999) (Australia)
K 5.142–6.215 4.1–12.0
Na 0.0319–0.0501 0.1–1.1
P 2.8–4.9
Mg 1.739–2.159 0.96–1.6
Ca 0.502–0.967 1.2–2.5
Cr 0.0113–0.0176
Mn 1.675–4.095 0.019–3.800
Fe 0.078–0.093 0.020–0.042
Co 0.0162–0.0166 0.0003
Ni 0.0120–0.0156
Cu 0.0048–0.0099 0.0031–0.0081
Zn 0.0403–0.0536 0.0021–0.0038
Pb 0.0108–0.0164
Cd not detected
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Australia and New Zealand, an acceptable alkaloid
dose for human consumption is up to 0.035 mg/kg/day
(Arnoldi & Greco 2011). White lupin lines of low
alkaloid content are characterised by a higher lysine
content than high-alkaloid lines (Green & Oram
1983). According to Annicchiarico et al. (2014),
in the subcontinental climate, the alkaloid content
in white lupin seeds may even be by 85% higher than
in the Mediterranean climate. White lupin alkaloids
present in seeds are also used for the control of the
occurrence of skin parasites in humans (Yeheyis et
al. 2010). In South America, bitter seeds of white
lupin are commonly used in medicine to reduce the
content of uronic acid and cholesterol in the blood
and fibre from white lupin seeds is used in a balanced
human diet (Torterolo et al. 2011).
Oligosaccharides. Seeds of sweet lines of white
lupin from California contained 5.3% of oligosac-
charides, including stachiosis (2.8%), sucrose (1.8%),
raffinose (0.4%), and verbascose (0.3%) (Mohamed
& rayas-duarte 1995), while in Australian cultivars
they constituted 5.85–7.41% in total (Petterson &
Fairbrother 1996). The content of oligosaccharides
depends on cultivar and also on the place of lupin
cultivation – more oligosaccharides were found in
Germany (10.3%) than in Argentina (8.6%) or Califor-
nia (5.3%) (mohamed & Rayas-duarte 1995). The
enzyme α-galactosidase, which is necessary for the
hydrolysis of α-1,6 bonds, is unavailable in the small
intestine of animals and humans. As a consequence,
these components pass to the large intestine, where
they undergo fermentation and gas production – CO2
,
methane, and H2
, giving well-known symptoms of
stomach ache and flatulence, from which people can
suffer after consuming a greater amount of legume
seeds (Petterson 1998). The anti-nutritive character
of soluble carbohydrates in lupin seeds also results
from their viscosity and their effect on bowel transit
time as well as on changes in hormone regulation and
different absorption of nutrients. In order to remove
them, seed germination is used, which is commonly
used in case of soybean (Enneking & Wink 2000).
It should be highlighted that recently these com-
pounds have been used more and more frequently for
the production of probiotics and prebiotics (Arnoldi
& Greco 2011). Oligosaccharides have a potential
value for the immune system, i.e. antioxidative ac-
tivity and antitumor activity, as well as lowering the
cholesterol level (Rochfort & Pannozo 2007). The
extraction of oligosaccharides from white lupin seeds
may affect a decrease in the content of γ-conglutin
and lipoxygenase (even by 37%), which however
may be beneficial due to their potential unfavour-
able allergenic effect, and the effect deteriorating
organoleptic traits of flour (Martinez-Villaluenga
et al. 2006). Oligosaccharide content in pasta pre-
pared with the use (10–30%) of white lupin flour was
significantly lower than the expected one resulting
from calculations, and after cooking it decreased by
30% (Lampart-Szczapa et al. 1997).
Inhibitors of protease. These are proteins which
strongly bind with digestive enzymes such as trypsin,
reduce their digestibility, and lead to malnutrition
or other disorders in animals. Inhibitors also have
a different function in plants, they are a source of
storage proteins which undergo breakdown at the
start of seed germination (Enneking & Wink 2000).
Inhibitors of trypsin may be deactivated through high
temperature treatment, which significantly improves
PER. White lupin seeds do not contain them at all
(they are mostly undetectable, i.e. less than 0.1 mg/g)
(Petterson & Fairbrother 1996).
Lectins. Also classified as glycoproteins, they are
capable of clumping red blood cells and they prevent
the absorption of nutrients. Lupin seeds contain small
amounts of lectins (3 × 10–5
HU, i.e. hemaglutina-
tion activity units) compared to e.g. common bean
(840 × 10–5
HU) (Nachbar & Oppenheim 1980) or
they are not detectable at all (Petterson & Fair-
brother 1996).
Phytic acid. It may reduce the bioavailability of
mineral components in monogastric animals through
cation chelation (Zn, Mg, Ca, Fe, K, and Mg) to non-
absorbable phytynians (Petterson & Fairbrother
1996; Petterson 1998). It should be highlighted that
phytic acid content in white lupin seeds (0.03%) is
significantly lower than e.g. in soybean – from 1.54%
(Mohamed & Rayas-Duarte 1995) to 1.89–2.27%
(Saastomoinen et al. 2013). In the studies of Mar-
tinez-Villaluenga et al. (2006) 0.025–0.044% of
phytic acid was found in white lupin seeds, while
according to Saastomoinen et al. (2013) – 0.063%.
Its content may be reduced even further through
fermentation or extrusion of seeds.
Phytochemicals
Tannins. Phenolic compounds produced at the end
of plant maturity and accumulated mainly in seed
coats constitute natural protection against diseases
and pests, and show antioxidative, antifungal, and an-
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tibacterial properties; however, Khan et al. (2015) did
not show any correlation between antioxidant activity
and phenolic contents. Petterson & Fairbrother
(1996) found small amounts of them in white lupin
seeds (0.23–0.52%), while Lampart-Szczapa et al.
(2009) detected slightly above 2 mg/100 g in dehulled
raw seeds. During seed fermentation, tannin con-
tent increased significantly up to about 8 mg/100 g,
whereas after extrusion it slightly decreased.
Isoflavones. They have a protective function in
plants and seeds and are synthesised in response to
biotic and abiotic stresses (phytoalexins). They have
been recognised as useful substances preventing the
occurrence of breast cancer, osteoporosis, and hot
flashes during menopause, they show hypocholes-
terolaemic properties (Khan et al. 2015), whereas
cholesterol content reduction is connected with
the stimulation of LDL fraction receptors (Sirtori
et al. 2004), most likely due to the reabsorption of
bile acids, which affects reduction in solubility and
in LDL cholesterol fractions (Martins & Bento
2007). The highest content of isoflavones in seeds
is characteristic of soybean. All green parts of white
lupin plants contain these compounds, and in seeds
(in the cell walls) their synthesis begins only in the
germination stage, while dry seeds are almost entirely
free of them (Katagiri et al. 2000).
Saponins. They are glycosides found in many plants
giving them bitter taste. White lupin seeds do not con-
tain them at all (Petterson & Fairbrother 1996)
or they were not detected (Muzquiz et al. 1993),
as distinguished from yellow lupin seeds (55.0 to
68.3 mg/kg), especially blue lupin (270.1–469.5 mg/kg)
(Muzquiz et al. 1993), and even 480–730 mg/kg in
Australian cultivars (Ruiz et al. 1995). In pea seeds,
their content is 1800 mg/kg, while in soybean seeds
up to 3500 mg/kg (Allen 1998). Saponins produce
complex compounds with bile acids and cholesterol
catching and removing their excess from the body.
Antioxidants. White lupin seeds in Saudi Arabia are
characterised by the highest content of tocopherols –
63.6 mg/100 g (α – 4.96 mg/100 g, δ – 4.10 mg/100 g,
and γ – 54.5 mg/100 g, and β fraction was not detect-
ed) (Alamri 2012), while in Australia 0.19 mg/100 g,
0.25 mg/100 g, and 20.1 mg/100 g, respectively (Frias
et al. 2005). The total amount of tocopherols in Eu-
rope according to Annicchiarico et al. (2014) was
12.1–13.3 g/100 g. In food products enriched with
lupin flour, tocopherols protect against free radi-
cals through inhibiting the oxidation of body lipids
and fats in the food. However, the cooking of white
lupin seeds causes a significant loss of tocopherols
(Kalogeropoulos et al. 2010).
Forms of white lupin seeds used
in the human diet
In the literature, information can be found on the
sensitivity of some people to lupin products (seeds,
flour, and dust) manifesting itself for example in
an anaphylactic shock in some individuals (Rad-
cliffe et al. 2005), urticaria, asthma, conjunctivitis,
swelling, or allergy of the mouth cavity (Sanz et al.
2010) as well as bloating, diarrhoea, and blurred
vision mainly in children after the consumption of
raw or roasted bitter white lupin seeds (Yeheyis et
al. 2010). Allergens are the main cause of allergic
reactions: Lup-1, which is a conglutin b (vicilin-like
protein) and Lup-2 corresponding to the conglutin
α fraction (legumin-like protein) (Guillamon et al.
2010). The problem of the allergenic effect of alka-
loids in white lupin seeds does not practically exist
now due to biological development and obtaining
sweet cultivars. The latest research has also indicated
that allergenic properties of lupin are significantly
lower than those of other leguminous species, e.g.
of soybean or peanuts, and concern only a small
percentage of the human population. However, the
European Commission included white lupin in the
list of allergens (Arnoldi & Greco 2011).
Nevertheless, white lupin seeds and their constitu-
ents used in different ways become an important mean
to meet or improve the health status and quality of
life not only in developing countries (Martirosyan
& Singh 2015). In Europe, mainly the seeds of white
lupin and partly blue lupin are used for consumption
purposes; the latter is commonly used in Australia.
White lupin was accepted for consumption purposes
by the Australian government in 1987 and by the
British one in 1996 (Swan 2000).
Sweethingham and Kingwell (2008) indicated
the most important areas of the use of lupin by hu-
mans, including the white lupin: whole seeds or
those without seed coats used in the traditional
fermentation of food in Asia, gluten-free lupin flour,
and protein fractions in the production of milk with-
out saturated fat, noodle and pasta products, as
substitutes of meat, egg protein, and sausages, etc.
Cooked, roasted, and ground into powder seeds are
also mixed with cereal flour in bread, crisps, and
pasta (Erbas et al. 2005). Roasted seeds may be used
102
Review Czech J. Food Sci., 35, 2017 (2): 95–105
doi: 10.17221/114/2016-CJFS
as snacks in the same way as e.g. peanuts. In roasted
and cooked seeds, alkaloid content is significantly
lower, and their dehulling decreases the content of
anti-nutritive compounds (Getachew et al. 2012).
Kalogeropoulos et al. (2010) observed a significant
decrease in total phenolic contents during cooking.
The addition of lupin flour to wheat flour allows for
obtaining more nutritive bread as far as protein and
dietary fibre is concerned. Mixing wheat and lupin
flour may help obtain the proper amino acid balance,
giving as a result a more complete food (Petterson
& Fairbrother 1996). Due to the better distribution
of water in dough, the obtained products have better
rheological properties, including better resistance
to freezing and thawing, the preparation of bread
dough can be easier, and shrinking can be limited.
White lupin seeds are also a perfect substrate for the
production of both bacterial and fungal fermented
products, e.g. tempe, miso, tofu, traditional sauces,
as well as dairy products (milk or yoghurt probiotics
similar to those produced from soybean seeds) in Asia
(Petterson & Fairbrother 1996; Petterson 1998;
Swan 2000) and Australia (Uauy et al. 1995). Lupin
seeds subjected to extrusion mixed with cereals are
also used for the production of crisps and dietary
dishes (Kiczorowska & Lipiec 2002). White lupin
sprouts contain about 41% protein and 8% fat, and
are a perfect potential source of food (Bhardwaj &
Hamama 2012a). From 1 kg of seeds you can obtain
even 7.5 kg of fresh sprout weight, and the sprouts
are practically free of lateral roots as distinguished
from e.g. soybean sprouts. The sprouts are also rich
in isoflavones as well as macro- and microelements,
especially Mn (139 mg/kg dry matter), and phytos-
terols increasing the nutritional value of white lupin.
Also germination is used as a process to increase
the nutritional value of many legumes (Khan et al.
2015). Seed germination causes a reduction in the
content of alkaloids, phytynians, and oligosaccharides
(Dagnia et al. 1992). On the other hand, the extru-
sion of white lupin seeds decreases the content of
fibre and crude fat, and slightly the content of most
amino acids (mainly methionine and cysteine) and
EAA value (Kiczorowska & Lipiec 2002). Owing
to the increased breakdown of proteins and com-
plex carbohydrates into simple and more digestible
structures of white lupin seeds, they were proved to
be better for fermentation than soybean seeds. The
completely new food applications of white lupin
include the use of immature seeds like in the case
of green peas or soybean (Bhardwaj & Hamama
2012b). The yield of immature white lupin pods of
a French cultivar Ludet was over 18 t/ha with the
protein content of 33%. Fermented white lupin seeds
for tinned fruit and vegetables are used in order to
increase pectin content (Golovchenko 2008), which
contain 4.5–5.5 g/100 g of the product, e.g. much
more than those obtained with other additives (e.g.
apple or beets) (Yeheyis et al. 2010).
The use of white lupin seeds by food companies
in Europe and Australia is, however, significantly
reduced, among other things, due to uncertainty of
the continuity of supplies of a sufficient amount and
quality of seeds, reduced access to knowledge con-
cerning their processing, and the risk of occurrence
of allergic reactions in a certain population (Uauy et
al. 1995; Sweetingham & Kingwell 2008).
Conclusion
White lupin is the oldest crop species of the kind
and may constitute a potential source of protein
without gluten. Despite valuable properties and
wide use, white lupin rather does not have a poten-
tial of being a global species. However, in the last
20 years, numerous properties of white lupin have
been discovered for the application in the produc-
tion of different kinds of functional food due to the
properties of seed components, which makes white
lupin seeds, regarding nutrition standards, more
valuable than other species of legumes.
The high biological value of proteins and important
sources of amino acids positively affect the physi-
ological condition of the human body, especially of
those suffering from diabetes, hypertension, obesity,
and cardiovascular diseases; they may also control
glucose content in diabetics or pre-diabetics, and
are especially suitable for vegans or for those on an
ovo-lacto vegetarian diet, or on a gluten-free diet
for people with the celiac disease. Human diet with
high lupin proteins affects a significant decrease in
serum cholesterol, including the LDL level, and it
also lowers the blood pressure. Lupin oil has desirable
ratios of omega-6 to omega-3 acids for consumption
purposes. White lupin seeds are also a valuable source
of dietary fibre, both soluble and insoluble, with its
well-known beneficial effect on reducing the cho-
lesterol level and on susceptibility to the ischaemic
heart disease. Food containing white lupin may also
suppress appetite, control glycaemia, improve the
lipid concentration in blood, and prevent colorectal
103
Czech J. Food Sci., 35, 2017 (2): 95–105 Review
doi: 10.17221/114/2016-CJFS
cancer; a low glycaemic index can prevent diseases
related to insulin resistance.
Part of specific substances contained in seeds may
have an unwanted effect (possibly the excessive alkaloid
content), but many of them have the potential value for
the immune system (e.g. oligosaccharides with their
antioxidative and antitumor activity, or lowering the
cholesterol content), others in turn occur in minimum
amounts (inhibitors of proteases, saponins), or there
is significantly less of them (phytic acid, lectins, iso-
flavones) than e.g. in soybean seeds. The problem of
allergenic effects of white lupin seeds concerns only
some kinds of protein, and much less alkaloids.
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Accepted after corrections: 2017–01–25
Published online: 2017–04–07

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White Lupin (Lupinus albus L.) – Nutritional and Health Values in Human Nutrition – a Review

  • 1. 95 Czech J. Food Sci., 35, 2017 (2): 95–105 Review doi: 10.17221/114/2016-CJFS White Lupin (Lupinus albus L.) – Nutritional and Health Values in Human Nutrition – a Review Janusz Prusinski* Department of Agrotechnology, UTP University of Science and Technology, Bydgoszcz, Poland *Corresponding author: [email protected] Abstract Prusinski J. (2017): White lupin (Lupinus albus L.) – nutritional and health values in human nutrition – a review. Czech J. Food Sci., 35: 95–105. White lupin seeds have been used in human nutrition and treatment for several thousand years. Nowadays the use of white lupin seeds is limited by a small scale of their production. However, in the last 20 years quite new properties of white lupin have been discovered for the application in the production of different kinds of functional food. Unique traits of protein, fatty acids with a desirable ratio of omega-6 to omega-3 acids, and fibre as well as other specific components, for example oligosaccharides and antioxidants or non-starch carbohydrates, make white lupin an excel- lent component in many healthy diets. The effects of white lupin components concern the physiological condition of the human body, including diabetes, hypertension, obesity, cardiovascular diseases, lipid concentration, glycaemia, appetite, insulin resistance, and colorectal cancer. Seeds are used among others for the production of gluten-free flour, bacterial and fungal fermented products, noodle and pasta products, as substitutes of meat, egg protein and sausages, also are cooked, roasted and ground and mixed with cereal flour in the production of bread, crisps and pasta, crisps and dietary dishes. Keywords: health benefits; chemical composition; effects on human health Many researchers state that the use of lupin seeds for consumption and medicinal purposes has been the subject of interest for more than 3000 years around the Mediterranean Sea. White lupin is the longest known crop species in the history of the genus Lupinus – it was known among Aegean farm- ers at least 400 years B.C. Until the beginning of the 19th century in Europe – in the Mediterranean Sea region it was the most often cultivated lupin species for green manure and for seeds, which were used in animal and human nutrition. However, because of high alkaloid content, the seeds could not be con- sidered as a safe food component (Prusinski 2015). A traditional way to avoid bitter taste caused by the presence of alkaloids in the lupin seeds was their very fine grinding, and next multiple rinsing with water, which however caused a decrease in the nutritive value as a result of removing soluble proteins, free amino acids, carbohydrates, and minerals. Neverthe- less, this method was commonly used by societies in Mediterranean Europe and Andean countries (Petterson 1998). After a significant success in qualitative cultiva- tion in the 20th century (Prusinski 2015) a signifi- cantly increased interest was observed concerning white lupin among feed and food producers, and also, among others, in medicine, food processing, and even in cosmetic industry, in the production of ecological pesticides, etc. Laboratory tests were very promising, however, the commercial use of white lupin seeds for the above-mentioned purposes is still insufficient and limited (Swan 2000). Meanwhile, due to the development of lifestyle diseases caused by an improper diet, i.e. cardiovascular diseases,
  • 2. 96 Review Czech J. Food Sci., 35, 2017 (2): 95–105 doi: 10.17221/114/2016-CJFS obesity, and diabetes, special attention is paid to healthy nutrition. White lupin seeds provide numer- ous health benefits counteracting the problems which are also connected with high blood pressure, insulin resistance, or higher cholesterol level. Taking into consideration predictions concerning a continuously growing number of deaths as a result of lifestyle diseases, functional food, including the use of lupin seeds, will have a greater and greater importance (Martins & Bento 2007; Martirosyan & Singh 2015). In many countries of Europe, the conservative attitude of consumers towards genetically modified organisms inhibited work on the popularisation of GMO in a significant way, and caused aversion towards importing products obtained from GMO cultivars (Sweetingham & Kingwell 2008). In the future, the lack of GMO lupin may lead to a sig- nificant development of a functionally varied food, mainly from white lupin seeds. Basic seed composition and properties of its constituents White lupin may be cultivated on almost all conti- nents, however, chemical composition of seeds and their nutritive value depend on plant response to environmental conditions. Only the protein content does not significantly depend on the place of cultiva- tion, while the proportion of other components in seeds is significantly dependent on the cultivation region (Bhardwaj et al. 1998). Protein. Owing to symbiosis with rhizobia, white lupin has an ability to fix N2 with the use of atmo- spheric nitrogen for the production of protein and other nitrogenous substances in seeds, which contain hardly any starch at the same time (Kurlovich et al. 2002). The average protein content in the seeds of white lupin is from 32.9% (Martinez-Villaluenga et al. 2006; Strakova et al. 2006) up to more than 36.0% (Sujak et al. 2006), or even 38.0% (Vecerek et al. 2008; Saastomoinen et al. 2013). Storage proteins in lupin consist of 85% globulins and 15% albumins (Petterson 1998). The globulin frac- tion contains 3 main proteins of varied amino acid composition: α-, ß-, and γ-conglutin. γ-Conglutin containing more methionine, cysteine, and valine is a sulphur-containing amino acid that constitutes about 4% of the protein composition (Duranti et al. 1981). Some of them may have an allergenic effect (Guillamon et al. 2010), however, compared to other legumes (peas, soybean, bean), white lupin seeds have a minimum content of proteins with anti-nutritive properties (Kurlovich et al. 2002) and a higher content of arginine, lysine, leucine, and phenylalanine than for example soybean, which makes white lupin seeds more valuable than other species regarding nutrition standards (Table 1). White lupin contains more amino acids (AA), including essential amino acids (EAA), and is also characterised by a higher index of essential amino acids (EAAI) and protein efficiency ratio (PER), as well as a higher nutritive value of protein isolates (CS) than the other two lupin species (yellow and narrow-leafed) cultivated in Europe (Sujak et al. 2006). Such composition indicates a high suitability of white lupin protein for vegans or for those on ovo-lacto vegetarian diet or on gluten-free diet for people with the celiac disease (Arnoldi & Greco 2011). Dehulling of white lupin seeds increases crude protein content and contents of most amino acids (except alanine) (Straková et al. 2006) leading to an increase of their biological value both for human nutrition and for animal feeding. On the other hand, extrusion of seeds leads to a decrease of amino acid content and the value of EAA (Kiczorowska & Lipiec 2002). High-protein diet may play an important role in ageing processes, pregnancy and lactation, growth of youth and athletes, as well as in recovery. Some white lupin proteins or their parts per se may posi- tively affect the physiological condition of human body under a wide range of unfavourable conditions, including diabetes, hypertension, obesity, cardiovas- cular diseases, or they may control glucose content in diabetics or pre-diabetics (Duranti 2011). Lupin flour contains approximately 30–40% protein, and effectiveness of its utilization is slightly lower than that of animal protein, though still satisfactory. The human diet which is high in lupin proteins affects a significant decrease in serum cholesterol, including the LDL (low-density lipoprotein, bad cholesterol) level, and also the level of triglycerides and glucose, and it also lowers the blood pressure (Arnoldi 2005; Nowicka et al. 2006). According to Naruszewicz et al. (2006), thanks to white lupin addition to food, a decrease in glucose content in the blood was 6.7%, homocysteine 11.8%, and high-sensitivity C-reactive protein (Hs-CRP) up to 18.3%. The favourable effect of soybean protein is already known, it can systematically decrease cholesterol
  • 3. 97 Czech J. Food Sci., 35, 2017 (2): 95–105 Review doi: 10.17221/114/2016-CJFS content in humans and animals, and also reduce the risk of the ischaemic heart disease. However, modern industry is searching for products of specific properties: technological and sensory ones, without phytoestrogens and affordably priced. Clinical tests proved capabilities of protein isolated from white lupin to reduce cholesterol content to the same ex- tent as it is observed in the case of other legumes (Sirtori et al. 2004). The absence of phytoestrogens in white lupin seeds makes this species better for the young (Sirtori & Naruszewicz 2005). Fat. Total fat content in white lupin seeds is from about 8% (Straková et al. 2006; Uzun et al. 2007; Vecerek et al. 2008; Andrzejewska et al. 2016) to 11.5% (Sujak et al. 2006). In the subcontinen- tal climate white lupin seeds are characterised by an approximately 8% lower fat content than in the Mediterranean climate (Annicchiarico et al. 2014). White lupin seeds are an interesting source of favourable ratios of important fatty acids used in the prophylaxis of circulatory system diseases (Si- mopoulos 2003). In their development, people have consumed fats in which the ratio of ω-6 to ω-3 acids was similar, whereas now in the diet of highly-developed countries it is up to 15 to 1. Excess ω-6 acids in the diet constitute a risk factor, while the 2 : 1 ratio, which is observed in white lupin seeds, has an enormous effect and decreases mortality as- sociated with circulatory system diseases. According to Green and Oram (1983), Arnoldi and Greco (2011), and Andrzejewska et al. (2016) oleic acid (mostly > 50%) prevails in the fat of white lupin seeds, which indicates a high suitability of lupin oil for consumption purposes (Table 2). In the seeds of bitter white lupin cultivars (spp. termis) the pro- portion of oleic acid is significantly higher (52.2 ± Table 1. Amino acid composition of white lupin seeds (g/16 g N) Specification Sujak et al. (2006) (Europe) Petterson & Fairbrother (1996) (Australia) Bhardwaj (2002) (USA) Essential amino acids Lysine 4.9 4.31 1.19 Methionine + cysteine 2.5 1.96 Cysteine 1.9 0.63 Threonine 3.5 3.16 0.83 Isoleucine 4.3 3.69 1.26 Tryptophan 0.6 0.97 Valine 4.1 3.55 1.06 Leucine 7.8 6.05 1.92 Histidine 3.3 1.74 0.51 Phenylalanine 5.6 3.33 1.02 Phenylalanine + tyrosine 5.6 8.68 Tyrosine 1.7 1.32 Non-essential amino acids Arginine 11.4 2.98 Asparagine 10.5 2.96 Serine 4.5 0.99 Glutamine 23.5 6.08 Proline 3.5 1.31 Glycine 4.3 0.92 Alanine 3.2 0.83 Nutritional values Amino acids participation (g/16 g N) 97.7 Essential amino acid participation (g/16 g N) 33.5 Chemical score of restrictive amino acid(s) 73.8 Essential amino acids index 85.0 Protein efficiency ratio 2.87
  • 4. 98 Review Czech J. Food Sci., 35, 2017 (2): 95–105 doi: 10.17221/114/2016-CJFS 2.32%) than in sweet cultivars (spp. albus) (44.9 ± 2.16%) (Alamri 2012). Andrzejewska et al. (2016) identified in white lupin seeds also the following ac- ids: myristic, pentadecanoic, palmitoleic, margaric, arachidic, eicosadienoic, eicosatrienoic, docosadi- enoic, and lignoceric at a content not higher than 1% each. Fatty acid forms (S) in Turkey (Erbas et al. 2005) were similar to those obtained by Bhardwaj (2002) in the USA. Carbohydrates. Lupin seeds and seed coats con- tain various types of carbohydrates, mainly non- starch carbohydrates (Khan et al. 2015), which are the most abundant in seeds. In the seed coat, apart from a small amount of proteins and fats, structural polysaccharides prevail: cellulose, hemicellulose, and pectins, while in the cotyledons – non-structural polysaccharides of cell walls (Straková et al. 2006; Vecerek et al. 2008), with the major proportion of galactose, arabinose, and uronic acid (Mohamed & Rayas-Duarte 1995; Petterson 1998). Most carbohydrates represent soluble or insoluble fibre up to about 2.83 g/100 g dry matter (Martinez- Villaluenga et al. 2006). Lupin starch is slowly digested and thus gradually releases glucose into the blood. However, according to some authors, mature air-dried white lupin seeds do not contain any starch (Petterson 1998; Borek et al. 2011) or its content is very low (Mohamed & Ryas Duarte 1995; Martinez-Villaluenga et al. 2006) (Table 3). As a result, lupin seeds indicate a low glycaemic index (Gullion & Champ 2002) and can prevent diseases related to insulin resistance. In Czech studies starch content in white lupin seeds was at least twice higher (Straková et al. 2006; Vecerek et al. 2008), which is however still very low compared to starch content e.g. in pea seeds (53.6–57.2%) (Dostálová et al. 2009). Crude fibre. White lupin seeds are a valuable source of dietary fibre (mostly insoluble), which is higher than in soybean seeds (Pisarikova & Zraly 2010). In most Fabaceae plants the content of crude fibre ranges from 8% to 27.5%, and that of soluble fibre from 3.3% to 13.8% (Guillon & Champ 2002). The average content of total fibre is from 101 g/kg (Tizazu & Emire 2010) to 367 g/kg (Martinez- Villaluenga et al. 2006) (Table 4). The seed coat in white lupin, which constitutes usually less than 20% of seed weight (Tizazu & Emire 2010), is mainly composed of cellulose with a small proportion of lignins. According to Pet- Table 2. Triglyceride composition in the seeds of white lupin (%) Fatty acids Andrzejewska et al. (2015) (Europe) Alamri (2010) (Asia) Bhardwaj (2002) (USA) Oleic acid (18:1) (n-9) 54.3 44.9 50.9 Linoleic acid (18:2) (n-6) 14.9 26.2 23.5 Palmitic acid (16:0) 8.57 7.71 6.60 Linolenic acid (18:3) (n-3) 7.22 15.8 9.68 Gadoleic acid (C20:1) (n-9) 4.14 Erucic acid 1.59 Stearic acid (18:0) 1.57 1.71 0.78 Arachidic acids (20:0) and erucic acid (22:1) 0.81 + 1.59 2.74 Σ SFA (saturated fatty acids) 16.1 9.63 11.3 Σ MUFA (monosaturated fatty acids) 58.8 55.4 Σ PUFA (polyunsaturated fatty acids) 15.0 9.38 33.3 Total n-6 15.0 Total n-3 7.22 n-6/n-3 2.11 Table 3. Content of starch in white lupin seeds (g/kg) Source Content Mohamed & Ryas Duarte (1995) 30.0 Petterson (1998) not detected Strakova et al. (2006) 81.5–85.7 Martinez-Villaluenga et al. (2006) 30.4 Vecerek et al. (2008) 72.6–86.5 Borek et al. (2011) not detected
  • 5. 99 Czech J. Food Sci., 35, 2017 (2): 95–105 Review doi: 10.17221/114/2016-CJFS terson (1998), there is up to 95 g of fibre in 100 g of the seed coat of white lupin. Thus, the dehulling of white lupin seeds decreases the total fibre con- tent and increases the soluble dietary fibre content (Pisarikova & Zraly 2010). Fibre contained in the cotyledons constituting about 30–40% of the germ weight consists of insoluble materials of the cell walls of a structure similar to pectins, which are soluble fibre known for its favourable effect on lowering the total cholesterol level as well as reducing susceptibil- ity to the ischaemic heart disease (Hall et al. 2005). Ash. White lupin seeds are a rich source of macro and microelements (Table 5); their total content is 30–40 mg/kg (Straková et al. 2006; Sujak et al. 2006; Saastamoinen et al. 2013). Among macroele- ments K, Mn, and Mg definitely prevail and among microelements Ca, Fe, and Na are dominant (Ka & Chandravanshi 2014). Lower Ca content improves culinary properties of seeds and reduces the time of their overcooking (Tizazu & Emire 2010). Compounds of favourable/unfavourable effect on human health Anti-nutritional compounds. The negative effect of unfavourable compounds in white lupin seeds which is often cited in literature is outdated to a significant extent now. As distinguished from other legumes (peas, soybean, and bean), white lupin seeds are characterised by a low or very low content of anti- nutritive substances (Muzquiz et al. 1998; Enne- king & Wink 2000). Their removal is possible either through selection of genotypes with a low content of these components or through post-harvest treatments, e.g. germinating, cooking, soaking, fermentation, ex- traction, etc. Quinolizidine alkaloids (QAs) are a family of about 100 bitter components – secondary metabolites of a bicyclic, tricyclic, and tetracyclic structure (Pet- terson 1998). Seeds of wild lupin species may even contain over 10 000 mg/kg alkaloids. The primary role of alkaloids was to protect plants against herbivorous animals. In their development, the alkaloid concen- tration in different plant parts (leaves, roots, stems) changes, reaching the highest value in the flowering stage. In Australian studies on white lupin the oc- currence of 4 alkaloids was observed: albine (15%), 13-hydroxylupanine (8%), lupanine (70%), and mul- tiflorine (3%) in the total amount not reaching over 0.01% (Petterson & Fairbrother 1996), while in Spanish studies additional a-isolupanine (Muzquiz et al. 1994) as well as 13-α-angeloyloxylupanine and 13-tigloyloxylupanine were reported (Muzquiz et al. 1998). Total alkaloid content in sweet white lu- pin cultivars has been significantly reduced in the process of domestication and breeding and does not currently exceed 0.02% (Prusinski 2015). Most alkaloids may cause cramps, vomiting, and even death as a result of the respiratory system pa- ralysis. Alkaloids also negatively affect the central nervous system in mammals, though in very low doses they may have a stimulating effect, while in high doses an inhibiting effect (McKnickiene & Asakaqviciute 2008). In humans too high dos- age, especially of lupanine and sparteine, may also cause trembling, arousal, and convulsions leading to blurred vision, dry mouth, nervousness, and bad mood (Arnoldi & Greco 2011). In many countries, among others in France, Great Britain, Australia, and New Zealand, the maximum alkaloid content in flour and lupin products was established on the level up to 200 mg/kg seeds (Resta et al. 2008). In Table 4. Content of crude fibre in white lupin seeds (g/kg) Source Content Martinez-Villaluenga et al. (2006) 367 Sujak et al. (2006) 144 Straková et al. (2006) 107 Vecerek et al. (2008) 134 Tizazu & Emire (2010) 101 Table 5. Content of macro and minor elements of ash in white lupin seeds (g/kg) Element Ka & Chandravanshi (2014) (Africa) Petterson & Fairbrother (1999) (Australia) K 5.142–6.215 4.1–12.0 Na 0.0319–0.0501 0.1–1.1 P 2.8–4.9 Mg 1.739–2.159 0.96–1.6 Ca 0.502–0.967 1.2–2.5 Cr 0.0113–0.0176 Mn 1.675–4.095 0.019–3.800 Fe 0.078–0.093 0.020–0.042 Co 0.0162–0.0166 0.0003 Ni 0.0120–0.0156 Cu 0.0048–0.0099 0.0031–0.0081 Zn 0.0403–0.0536 0.0021–0.0038 Pb 0.0108–0.0164 Cd not detected
  • 6. 100 Review Czech J. Food Sci., 35, 2017 (2): 95–105 doi: 10.17221/114/2016-CJFS Australia and New Zealand, an acceptable alkaloid dose for human consumption is up to 0.035 mg/kg/day (Arnoldi & Greco 2011). White lupin lines of low alkaloid content are characterised by a higher lysine content than high-alkaloid lines (Green & Oram 1983). According to Annicchiarico et al. (2014), in the subcontinental climate, the alkaloid content in white lupin seeds may even be by 85% higher than in the Mediterranean climate. White lupin alkaloids present in seeds are also used for the control of the occurrence of skin parasites in humans (Yeheyis et al. 2010). In South America, bitter seeds of white lupin are commonly used in medicine to reduce the content of uronic acid and cholesterol in the blood and fibre from white lupin seeds is used in a balanced human diet (Torterolo et al. 2011). Oligosaccharides. Seeds of sweet lines of white lupin from California contained 5.3% of oligosac- charides, including stachiosis (2.8%), sucrose (1.8%), raffinose (0.4%), and verbascose (0.3%) (Mohamed & rayas-duarte 1995), while in Australian cultivars they constituted 5.85–7.41% in total (Petterson & Fairbrother 1996). The content of oligosaccharides depends on cultivar and also on the place of lupin cultivation – more oligosaccharides were found in Germany (10.3%) than in Argentina (8.6%) or Califor- nia (5.3%) (mohamed & Rayas-duarte 1995). The enzyme α-galactosidase, which is necessary for the hydrolysis of α-1,6 bonds, is unavailable in the small intestine of animals and humans. As a consequence, these components pass to the large intestine, where they undergo fermentation and gas production – CO2 , methane, and H2 , giving well-known symptoms of stomach ache and flatulence, from which people can suffer after consuming a greater amount of legume seeds (Petterson 1998). The anti-nutritive character of soluble carbohydrates in lupin seeds also results from their viscosity and their effect on bowel transit time as well as on changes in hormone regulation and different absorption of nutrients. In order to remove them, seed germination is used, which is commonly used in case of soybean (Enneking & Wink 2000). It should be highlighted that recently these com- pounds have been used more and more frequently for the production of probiotics and prebiotics (Arnoldi & Greco 2011). Oligosaccharides have a potential value for the immune system, i.e. antioxidative ac- tivity and antitumor activity, as well as lowering the cholesterol level (Rochfort & Pannozo 2007). The extraction of oligosaccharides from white lupin seeds may affect a decrease in the content of γ-conglutin and lipoxygenase (even by 37%), which however may be beneficial due to their potential unfavour- able allergenic effect, and the effect deteriorating organoleptic traits of flour (Martinez-Villaluenga et al. 2006). Oligosaccharide content in pasta pre- pared with the use (10–30%) of white lupin flour was significantly lower than the expected one resulting from calculations, and after cooking it decreased by 30% (Lampart-Szczapa et al. 1997). Inhibitors of protease. These are proteins which strongly bind with digestive enzymes such as trypsin, reduce their digestibility, and lead to malnutrition or other disorders in animals. Inhibitors also have a different function in plants, they are a source of storage proteins which undergo breakdown at the start of seed germination (Enneking & Wink 2000). Inhibitors of trypsin may be deactivated through high temperature treatment, which significantly improves PER. White lupin seeds do not contain them at all (they are mostly undetectable, i.e. less than 0.1 mg/g) (Petterson & Fairbrother 1996). Lectins. Also classified as glycoproteins, they are capable of clumping red blood cells and they prevent the absorption of nutrients. Lupin seeds contain small amounts of lectins (3 × 10–5 HU, i.e. hemaglutina- tion activity units) compared to e.g. common bean (840 × 10–5 HU) (Nachbar & Oppenheim 1980) or they are not detectable at all (Petterson & Fair- brother 1996). Phytic acid. It may reduce the bioavailability of mineral components in monogastric animals through cation chelation (Zn, Mg, Ca, Fe, K, and Mg) to non- absorbable phytynians (Petterson & Fairbrother 1996; Petterson 1998). It should be highlighted that phytic acid content in white lupin seeds (0.03%) is significantly lower than e.g. in soybean – from 1.54% (Mohamed & Rayas-Duarte 1995) to 1.89–2.27% (Saastomoinen et al. 2013). In the studies of Mar- tinez-Villaluenga et al. (2006) 0.025–0.044% of phytic acid was found in white lupin seeds, while according to Saastomoinen et al. (2013) – 0.063%. Its content may be reduced even further through fermentation or extrusion of seeds. Phytochemicals Tannins. Phenolic compounds produced at the end of plant maturity and accumulated mainly in seed coats constitute natural protection against diseases and pests, and show antioxidative, antifungal, and an-
  • 7. 101 Czech J. Food Sci., 35, 2017 (2): 95–105 Review doi: 10.17221/114/2016-CJFS tibacterial properties; however, Khan et al. (2015) did not show any correlation between antioxidant activity and phenolic contents. Petterson & Fairbrother (1996) found small amounts of them in white lupin seeds (0.23–0.52%), while Lampart-Szczapa et al. (2009) detected slightly above 2 mg/100 g in dehulled raw seeds. During seed fermentation, tannin con- tent increased significantly up to about 8 mg/100 g, whereas after extrusion it slightly decreased. Isoflavones. They have a protective function in plants and seeds and are synthesised in response to biotic and abiotic stresses (phytoalexins). They have been recognised as useful substances preventing the occurrence of breast cancer, osteoporosis, and hot flashes during menopause, they show hypocholes- terolaemic properties (Khan et al. 2015), whereas cholesterol content reduction is connected with the stimulation of LDL fraction receptors (Sirtori et al. 2004), most likely due to the reabsorption of bile acids, which affects reduction in solubility and in LDL cholesterol fractions (Martins & Bento 2007). The highest content of isoflavones in seeds is characteristic of soybean. All green parts of white lupin plants contain these compounds, and in seeds (in the cell walls) their synthesis begins only in the germination stage, while dry seeds are almost entirely free of them (Katagiri et al. 2000). Saponins. They are glycosides found in many plants giving them bitter taste. White lupin seeds do not con- tain them at all (Petterson & Fairbrother 1996) or they were not detected (Muzquiz et al. 1993), as distinguished from yellow lupin seeds (55.0 to 68.3 mg/kg), especially blue lupin (270.1–469.5 mg/kg) (Muzquiz et al. 1993), and even 480–730 mg/kg in Australian cultivars (Ruiz et al. 1995). In pea seeds, their content is 1800 mg/kg, while in soybean seeds up to 3500 mg/kg (Allen 1998). Saponins produce complex compounds with bile acids and cholesterol catching and removing their excess from the body. Antioxidants. White lupin seeds in Saudi Arabia are characterised by the highest content of tocopherols – 63.6 mg/100 g (α – 4.96 mg/100 g, δ – 4.10 mg/100 g, and γ – 54.5 mg/100 g, and β fraction was not detect- ed) (Alamri 2012), while in Australia 0.19 mg/100 g, 0.25 mg/100 g, and 20.1 mg/100 g, respectively (Frias et al. 2005). The total amount of tocopherols in Eu- rope according to Annicchiarico et al. (2014) was 12.1–13.3 g/100 g. In food products enriched with lupin flour, tocopherols protect against free radi- cals through inhibiting the oxidation of body lipids and fats in the food. However, the cooking of white lupin seeds causes a significant loss of tocopherols (Kalogeropoulos et al. 2010). Forms of white lupin seeds used in the human diet In the literature, information can be found on the sensitivity of some people to lupin products (seeds, flour, and dust) manifesting itself for example in an anaphylactic shock in some individuals (Rad- cliffe et al. 2005), urticaria, asthma, conjunctivitis, swelling, or allergy of the mouth cavity (Sanz et al. 2010) as well as bloating, diarrhoea, and blurred vision mainly in children after the consumption of raw or roasted bitter white lupin seeds (Yeheyis et al. 2010). Allergens are the main cause of allergic reactions: Lup-1, which is a conglutin b (vicilin-like protein) and Lup-2 corresponding to the conglutin α fraction (legumin-like protein) (Guillamon et al. 2010). The problem of the allergenic effect of alka- loids in white lupin seeds does not practically exist now due to biological development and obtaining sweet cultivars. The latest research has also indicated that allergenic properties of lupin are significantly lower than those of other leguminous species, e.g. of soybean or peanuts, and concern only a small percentage of the human population. However, the European Commission included white lupin in the list of allergens (Arnoldi & Greco 2011). Nevertheless, white lupin seeds and their constitu- ents used in different ways become an important mean to meet or improve the health status and quality of life not only in developing countries (Martirosyan & Singh 2015). In Europe, mainly the seeds of white lupin and partly blue lupin are used for consumption purposes; the latter is commonly used in Australia. White lupin was accepted for consumption purposes by the Australian government in 1987 and by the British one in 1996 (Swan 2000). Sweethingham and Kingwell (2008) indicated the most important areas of the use of lupin by hu- mans, including the white lupin: whole seeds or those without seed coats used in the traditional fermentation of food in Asia, gluten-free lupin flour, and protein fractions in the production of milk with- out saturated fat, noodle and pasta products, as substitutes of meat, egg protein, and sausages, etc. Cooked, roasted, and ground into powder seeds are also mixed with cereal flour in bread, crisps, and pasta (Erbas et al. 2005). Roasted seeds may be used
  • 8. 102 Review Czech J. Food Sci., 35, 2017 (2): 95–105 doi: 10.17221/114/2016-CJFS as snacks in the same way as e.g. peanuts. In roasted and cooked seeds, alkaloid content is significantly lower, and their dehulling decreases the content of anti-nutritive compounds (Getachew et al. 2012). Kalogeropoulos et al. (2010) observed a significant decrease in total phenolic contents during cooking. The addition of lupin flour to wheat flour allows for obtaining more nutritive bread as far as protein and dietary fibre is concerned. Mixing wheat and lupin flour may help obtain the proper amino acid balance, giving as a result a more complete food (Petterson & Fairbrother 1996). Due to the better distribution of water in dough, the obtained products have better rheological properties, including better resistance to freezing and thawing, the preparation of bread dough can be easier, and shrinking can be limited. White lupin seeds are also a perfect substrate for the production of both bacterial and fungal fermented products, e.g. tempe, miso, tofu, traditional sauces, as well as dairy products (milk or yoghurt probiotics similar to those produced from soybean seeds) in Asia (Petterson & Fairbrother 1996; Petterson 1998; Swan 2000) and Australia (Uauy et al. 1995). Lupin seeds subjected to extrusion mixed with cereals are also used for the production of crisps and dietary dishes (Kiczorowska & Lipiec 2002). White lupin sprouts contain about 41% protein and 8% fat, and are a perfect potential source of food (Bhardwaj & Hamama 2012a). From 1 kg of seeds you can obtain even 7.5 kg of fresh sprout weight, and the sprouts are practically free of lateral roots as distinguished from e.g. soybean sprouts. The sprouts are also rich in isoflavones as well as macro- and microelements, especially Mn (139 mg/kg dry matter), and phytos- terols increasing the nutritional value of white lupin. Also germination is used as a process to increase the nutritional value of many legumes (Khan et al. 2015). Seed germination causes a reduction in the content of alkaloids, phytynians, and oligosaccharides (Dagnia et al. 1992). On the other hand, the extru- sion of white lupin seeds decreases the content of fibre and crude fat, and slightly the content of most amino acids (mainly methionine and cysteine) and EAA value (Kiczorowska & Lipiec 2002). Owing to the increased breakdown of proteins and com- plex carbohydrates into simple and more digestible structures of white lupin seeds, they were proved to be better for fermentation than soybean seeds. The completely new food applications of white lupin include the use of immature seeds like in the case of green peas or soybean (Bhardwaj & Hamama 2012b). The yield of immature white lupin pods of a French cultivar Ludet was over 18 t/ha with the protein content of 33%. Fermented white lupin seeds for tinned fruit and vegetables are used in order to increase pectin content (Golovchenko 2008), which contain 4.5–5.5 g/100 g of the product, e.g. much more than those obtained with other additives (e.g. apple or beets) (Yeheyis et al. 2010). The use of white lupin seeds by food companies in Europe and Australia is, however, significantly reduced, among other things, due to uncertainty of the continuity of supplies of a sufficient amount and quality of seeds, reduced access to knowledge con- cerning their processing, and the risk of occurrence of allergic reactions in a certain population (Uauy et al. 1995; Sweetingham & Kingwell 2008). Conclusion White lupin is the oldest crop species of the kind and may constitute a potential source of protein without gluten. Despite valuable properties and wide use, white lupin rather does not have a poten- tial of being a global species. However, in the last 20 years, numerous properties of white lupin have been discovered for the application in the produc- tion of different kinds of functional food due to the properties of seed components, which makes white lupin seeds, regarding nutrition standards, more valuable than other species of legumes. The high biological value of proteins and important sources of amino acids positively affect the physi- ological condition of the human body, especially of those suffering from diabetes, hypertension, obesity, and cardiovascular diseases; they may also control glucose content in diabetics or pre-diabetics, and are especially suitable for vegans or for those on an ovo-lacto vegetarian diet, or on a gluten-free diet for people with the celiac disease. Human diet with high lupin proteins affects a significant decrease in serum cholesterol, including the LDL level, and it also lowers the blood pressure. Lupin oil has desirable ratios of omega-6 to omega-3 acids for consumption purposes. White lupin seeds are also a valuable source of dietary fibre, both soluble and insoluble, with its well-known beneficial effect on reducing the cho- lesterol level and on susceptibility to the ischaemic heart disease. Food containing white lupin may also suppress appetite, control glycaemia, improve the lipid concentration in blood, and prevent colorectal
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