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Designing selenium functional foods and beverages: A review
Accepted Manuscript Designing selenium functional foods and beverages: A review
Parise Adadi, Nadezhda V. Barakova, Kirill Y. Muravyov, Elena F. Krivoshapkina PII: DOI: Reference:
S0963-9969(18)30914-1 https://doi.org/10.1016/j.foodres.2018.11.029 FRIN 8092
To appear in:
Food Research International
Received date: Revised date: Accepted date:
23 July 2018 15 October 2018 15 November 2018
Please cite this article as: Parise Adadi, Nadezhda V. Barakova, Kirill Y. Muravyov, Elena F. Krivoshapkina , Designing selenium functional foods and beverages: A review. Frin (2018), https://doi.org/10.1016/j.foodres.2018.11.029
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ACCEPTED MANUSCRIPT Designing selenium functional foods and beverages: A review
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Parise Adadi*, Nadezhda V. Barakova, Kirill Y. Muravyov, Elena F. Krivoshapkina
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ITMO University, Lomonosova Street 9, 191002, St. Petersburg, Russian Federation
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*Corresponding author
ITMO University,
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Lomonosova Street 9, 191002,
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Parise Adadi
St. Petersburg, Russian Federation
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E-mail address: [email protected]; adadi_parise@scamt- itmo.ru.
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Tel: +79817511640
ACCEPTED MANUSCRIPT ABSTRACT A functional food is any food that has (a) specific nutrient(s) added to it for a specific functional purpose. Selenium (Se) is a metalloid that belongs to group 16 of the periodic table. It may be obtained from myriad sources like soil, water, and living organisms. Se is required to sustain proper health in both animals and humans due to its linkage with various biological functions in
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the immune system. Nature has made it impossible for us to obtain sufficient Se from the diet since some regions across the globe have been designated as Se-deficient while others are
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becoming Se-toxic. Se deficiency is associated with a compromised immune system and
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increased susceptibility to various diseases. Therefore, designing Se functional foods and supplements is the way forward in curbing the menace mentioned above, since geographical
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location will not have any effect on the Se content of these foods. Brewing yeast has the
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necessary enzymes to biotransform inorganic Se (Na 2 SeO 3 ) to its bioactive organic form, which is incorporated in the aged selenized beer. S. cerevisiae was found to have better
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biotransformation efficiency than other yeast species. A traditional Slavic beverage, selenized kvass, was brewed using rye grains soaked and germinated in solutions of Na 2 SeO 3 . Fruit yeasts
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(Lesaffre) and Evitalia were utilized as the starter cultures. A Se enriched solution was extracted from Se-biofortified pak-choi cabbage and incorporated into the wort before fermentation.
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Selenized yeast and algae can be taken as supplements or as food additives. There have been some reports about microcystins (MCs) in Spirulina. Therefore, the safety of Se-algae is not guaranteed. Pasteurized dried selenized supplements (yeast and algae) were proposed in formulating Selenized enriched Tom-brown. This review seeks to propose some possible foods that could be enriched with Se. A large portion of the population adequately consumes these proposed foods on a regular basis hence the target goal would be a success.
Keywords: Selenized beer, Slavic foods (Kvass), Yeast, Algae, Selenosis, Anti-cancer effects
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Abbreviations. FOSHU – foods for specified health use; FUFOSE – functional food science in Europe; SHF – specific health promoting food; TDI – tolerable daily intake; MCs – microcystins; SeCys – selenocysteine;
SeMeSeCys
–
selenomethyl-selenocysteine;
SeMet
–
selenomethionine;
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NaHSeO 3 – sodium hydroselenite; kDa – Kilo Dalton; Se-yeast – selenized-yeast; Se-algae – selenized algae; Se – Selenium; ICP-MS – liquid chromatography hyphenated to an inductively
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coupled plasma mass spectrometer Selenium; MHLW – Ministry of health, labor and welfare;
inductively
coupled
plasma
mass
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SOD – superoxide dismutase; CAT – catalase; GSH-Px – glutathione peroxidase; ICP-MS – spectrometry;
HPLC
–
High
Performance
Liquid
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Chromatography; MMeSe – monomethyl form of Se; DMeSe – dimethyl form of Se; TMeSe –
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trimethyl form of Se; SDS – Sodium dodecyl sulfate.
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1. Introduction In the last few decades, the world has experienced an increase in the cost of healthcare and people have found this alarming. Hence, they are searching for ways on how to stay healthy in order to avoid excessive spending on healthcare. Presently, the perception of food has changed
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from just consuming/drinking something to fill the stomach but rather eating to benefit from the nutrition, which enhances the physiological functioning of the system. At about this time the
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Japanese approved the process for functional foods called Foods for Specified Health Use
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(FOSHU) in the 1980s (Arai, 1996). There is no universal definition for functional foods as they vary across countries (The Japanese ([FOSHU]), European (Functional Food Science in Europe
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[FUFOSE]), and Dutch (Specific Health Promoting Food [SHF])). For instance, FUFOSE
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defined a functional food as “food that beneficially affects one or more target functions in the body, beyond adequate nutritional effects, in a way that is relevant to either an improved state of health and well-being and reduction of risk of disease. It is part of a normal food pattern. It is not
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a pill, a capsule or any form of dietary supplement.” The Academy of Nutrition and Dietetics
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sees it as “whole food along with fortified, enriched, or enhanced foods that have a potentially beneficial effect on health when consumed as part of a varied diet on a regular basis at effective
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levels.” Conversely, FOSHU described it as foods containing an ingredient with functions for health and officially approved to claim its physiological effects on the human body (Academy of Nutrition and Dietetics, 2013). Aryee & Boye (2015) have documented various health benefits of functional foods.
According to Cocks, Wrigley, Chicarelli-Robinson, & Smith (1995), herbs, algae, and microorganisms are rich sources of bioactive ingredients used in designing functional foods. Functional foods must remain foods (should not be pills or capsules) mostly composed of bulk ingredients (fruits, beverages, vegetables, cereals, nuts, milk, and milk-based products), and they
ACCEPTED MANUSCRIPT must demonstrate their effects in amounts that can be expected for normal consumption patterns (Sandhu & Sra, 2014). The demand for functional foods has increased over the past decades, resulting in the exploitation of this need by entrepreneurs who have established multibillion-dollar companies mainly invested in designing foods to satisfy consumers’ needs. These companies have seen
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significant growth, mostly in developed countries (United States, Canada, Denmark, Ireland, etc.).
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Selenium (Se) is a metalloid that belongs to group 16 of the Mendeleev's periodic table with
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physicochemical properties similar to sulfur due to their proximity (ionic radius) on the periodic table. The biological and toxicological effects of Se strongly depend on its chemical form,
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therefore the organic form is preferable to the inorganic form due to its bioavailability (Bodnar,
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Konieczka, & Namiesnik, 2012; Gupta & Gupta, 2016). Se is an essential nutrient in human life and participates in some biochemical reactions in the
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body. As reported by Oldfield (1999), Se is directly linked to a spectrum of biological activity. It exhibits similar antioxidant properties (act like superhero nutrients) to lycopene (Adadi,
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Barakova, & Krivoshapkina, 2018), thus protecting the body against damaging free radicals. Se act as a cofactor for triiodothyronine deiodinases, an important enzyme involved in thyroid
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hormone metabolism, thereby affecting iodine status and consequently preventing goiters (Arnaud et al., 2001; Erdenetsogt, Golubkina, Nadegkin, Monhoo, & Batjargal, 2014). It also acts as a cofactor for various enzymes, for example, glutathione peroxidase (GSH-Px), superoxide dismutase (SOD), and catalase (CAT), which shield the cells against damage triggered by free radicals and lipoperoxides. (Thompson & Scott, 1969; Newberne & Suphakarn, 1983). Health disorders such as infertility, cardiovascular diseases and cancer (oxidative stress) are associated with the insufficient intake of Se (Broadley et al., 2006). Human in vitro studies suggest that supplementing the diet with Se decreased the incidence of cancer (Combs, 2005; De Martino, Filomeni, Aquilano, Ciriolo, & Rotilio, 2006).
ACCEPTED MANUSCRIPT It is estimated that the diets of as many as 1 billion people might lack sufficient Se for their wellbeing (Combs, 2001; Fairweather-Tait, Collings, & Hurst, 2011; Joy et al., 2014; Stoffaneller & Morse, 2015). Nevertheless, Se deficiency and toxicity remain a problem in many human populations (National Research Council, 1983). Plant products are the major source of Se. Human low dietary Se intake is associated with crops
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cultivated in soils low in Se (Broadley et al., 2006; White & Broadley, 2009; Chilimba et al., 2011; Fairweather-Tait et al., 2011; Rayman, 2012; Fordyce, 2013; Joy et al., 2015).
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Consequently, animal-based food could serve as an alternate source since animals can
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accumulate Se from Se-supplemented feed or pastures fertilized with Se fertilizers. Meat, sausages, eggs, milk, apples (Se-biofortified apples), dairy products, fish are the primary sources
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of Se in the human diet, although this varies across regions (Meyer, Heerdegen, Schenkel,
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Dänicke, & Flachowsky, 2004; Rayman, 2008; Rayman, 2012; Wortmann, Enneking & Daum, 2018). Zhu, Pilon-Smits, Zhao, Williams, & Meharg (2009) found that some regions around the
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world are Se-deficient while others are becoming Se-toxic due to natural and anthropogenic activities.
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There is a direct correlation between soil Se content and the Se dose in products obtained from such soils (Finley, Matthys, Shuler, & Korynta, 1996). The soils in Sverdlovsk, Chelyabinsk,
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Tyumen, Novosibirsk, Irkutsk, Chita, Magadan, Amur, Krasnoyarsk, Khabarovsk, Primorsky Krai, Sakha-Yakutia, Komi, Mari-El Republic, Karelia, St. Petersburg, the Upper Volga region, Yaroslavl region and Udmurt Republic are naturally deficient in Se (Golubkina & Alfthan, 1999). This predisposes the inhabitants of these regions to mild selenosis with symptoms ranging from dermatitis, cracking of nails, hair loss and garlicky breath (due to exhalation of dimethylselenide). Severe selenosis can lead to acute respiratory distress, myocardial infarction and renal failure (White, 2015). As reported by Dhillon & Dhillon (2003) and Fordyce, (2013), animals also suffer from this disease, with symptoms ranging from
hair loss, garlicky breath,
hoof deformation (in horse, cattle, donkey), abnormal posture, lack of vitality, growth disorder,
ACCEPTED MANUSCRIPT anorexia, diarrhea, reduced reproductive performance, fetal deformities and respiratory failure due to insufficient intake of Se. Se deficiency is not only bound to Russia but other areas around the world (Zhu et al., 2009; Yuan et al., 2012). Se fertilizers were formulated to enable farmers to fertilize their crops (biofortification). Products from these farms were found to contain more Se than the control
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farms (Finley et al., 1996). The problem with toxicity and volatility of Se salts at high temperatures when applied also
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surface and remains a challenge to farmers. However, the problem with toxicity could be curbed
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by planting specialized crops, namely Se accumulators (e.g., medicinal melon) in the contaminated soil (White, 2015; Gupta & Gupta, 2016). For these reasons, Se-enriched foods
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and supplements have been proposed to circumvent the problems related to Se deficiency.
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Therefore, some technologies for designing Se-enriched functional foods, a mechanism for toxicities and health concern were discussed. The impact of Se on gut microbiota, its
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bioavailability, as well as the factors to consider when adopting the techniques mentioned above were also pondered in this manuscript.
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2. Production of beverages enriched with selenium Beverages are drinks prepared by the combination of two or more materials (i.e., water, sugar,
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colors, additives, yeast, etc.) for human consumption except for the regular water we drink every day. They are divided into two types (i) alcoholic (ii) non- alcoholic beverages. Beer, wine, and palm wine, etc., are categorized as alcoholic beverages while tea, kvass and fruit drinks are grouped under non-alcoholic beverages. These classifications vary according to the standard of each country. Beverages are not just for quenching thirst but also provide the consumers with energy, vitamins, minerals (Fellows & Hampton, 1992) and other vital bioactive substances required for the normal physiological function systems. In 2012, about 923 billion liters of commercialized beverages were sold across the world, with consumption in the United of States of America, Germany, and Brazil estimated at 182.2, 51.1,
ACCEPTED MANUSCRIPT 62.2 billion liters of beverages respectively in 2013 fiscal year alone (Euromonitor International, 2012; Madi, Castro, & Wallis, 2016). In producing these beverages, single or mixed starter cultures are usually employed. These cultures possess the vital enzymes to biotransform inorganic Se to more bioactive organic forms. As reported by Nelson, (2005), beer is one of the world’s most known and widely consumed drink after tea and water. Hence, enriching this
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product with Se could be an excellent intervention in combating deficiency of this micronutrient. While alcohol consumption is prohibited in Muslim countries, non-alcoholic beverages could
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also be designed as alternate conveyors to compensate the Se deficiencies in those countries.
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2.1 Supplementation of Se during brewing
Brewing yeast - Saccharomyces cerevisiae, and lactic acid bacteria (LAB) have shown the
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ability to transform inorganic Se to the organic form (safer, preferred and highly bioactive)
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(Schrauzer, 2000; Alzate et al., 2007; Alzate et al., 2008). This has drawn the attention of brewers to look into this novel technology in order to brew Se enriched beers.
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Metabolizing low bioavailable Se compounds (inorganic Se) into their high bioavailable forms (organic Se) by a plant, yeast, bacteria was reported (Kurek et al., 2016). Sodium selenite
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(Na2 SeO 3 ) was supplemented in media cultured with Bifidobacterium animalis (Zhang et al., 2009) as probiotic bacteria used in producing Se-enrich products. The single infusion method of
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mashing could be used, where the malt is mixed with distilled water in the mash tun and heated at 45 °C for 30 minutes. The temperature is then increased to 54 °C and then 62 °C with step times of 40 and 45 min respectively. After these step times, the temperature will once, again be increased to 72 °C for 55 min. Finally, the temperature is increased to 78 °C for 10 minutes. Filtration is carried out while 1 L distilled water is used for the sparging process. The wort is then boiled at 100 °C for 90 min, hops is then added 80 min before the end of the boiling time. The wort is then allowed to cool to the desired temperature (20 o C) by either placing the boiled wort under running cold water or using a chilled sterilized copper wort chiller. The wort is then transferred to sterilized fermentation vessels equipped with airlock bubblers (Adadi, Kovaleva,
ACCEPTED MANUSCRIPT Glukhareva, Shatunova, & Petrov, 2017; Adadi, Kovaleva, Glukhareva, & Barakova, 2018) where a different concentration of Na2 SeO 3 is added (Sánchez-Martínez et al., 2012). Even though Na2 SeO 3 could be used as a precursor for the formation of selenoproteins in beer, it cannot be stored for future use (Alfthan, Aro, Arvilommi, & Huttunen, 1991). The flow chart of brewing Se enriched beer is depicted in figure 1. The yeast is then pitched while the fermentation
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kinetics is studied. Fermentation is then carried out at a temperature of 25 °C for 5-12 days depending on the type of Se-enriched beer brewed.
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The total Se and Se species could be determined using inductively coupled plasma mass
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spectrometry (ICP-MS) and liquid chromatography hyphenated to an inductively coupled plasma mass spectrometer (HPLC–ICP-MS) respectively. The details are documented in Perez-Corona
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et al. (2011) and Sánchez-Martínez et al. (2012).
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The fermentation process is not altered due to the addition of Se in the wort or must (PerezCorona et al., 2011). Se does not alter the sugars and other nutrients vital for the yeast, hence the
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fermentation process will proceed without any problems. Yeasts are excellent accumulators of Se and other trace elements (Schrauzer, 2006; Kieliszek & 2013).
Saccharomyces
cerevisiae
and
Lactobacillus have the enzyme for
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Błażejak,
biotransformation of inorganic Se to organic form and they are widely utilized in brewing
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beverages (Sánchez-Martínez et al., 2012). Much research is devoted to the study of intracellular accumulation and metabolism of Se, mainly by yeast and bacteria (Ponce de León, Bayon, Paquin, & Caruso, 2002; Gharieb & Gadd 2004; Pieniz, Andreazza, Mann, Camargo, & Brandelli, 2017). Kieliszeket, Błażejak, Gientka, & Bzducha-Wróbe (2015) tried to decipher the accumulation and transformation of Se in yeast cells. The added Na2 SeO 3 is metabolized in two possible pathways, i.e. by methylation coupled with the reduction of Se and direct Se incorporation or bind to proteins in which Se replaces sulfur mainly by the amino acids cysteine and methionine (Fairweather-Tait et al., 2010; Kieliszek &
ACCEPTED MANUSCRIPT B1azejak, 2013). Kieliszek & B1azejak (2013), in their review, stated that selenates (SeO4 -2 ) are probably reduced to selenites (SeO 3 -2 ), elemental Se (Se0 ), and selenides (Se2-). In the presence of
yeast,
glutathione
and
adenosine-5-triphosphate
are
transformed
to
adenosine-5-
selenophosphate via enzymatic activation (i.e., Sul1 and Sul2 sulfate permeases, ATP sulfurylase, PAPSe reductase) in which SeO 4 -2 is converted to SeO 3 -2 . Other forms of Se such as
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monomethyl (monomethyl form of selenium (MMeSe), dimethyl (dimethyl form of Se (DMeSe), and trimethyl (trimethyl form of Se (TMeSe) can be obtained from non-methylated form, which
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is less toxic than other forms of Se (Pyrzynska, 1996; Bánszky, Simonics, & Maráz, 2003;
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Mapelli, Hillestrøm, Patil, Larsen, & Olsson, 2012; Kieliszek et al., 2015; Kieliszek & B1azejak, 2013). SeO 3 -2 reacts further with glutathione to form selenodiglutathione (GS-Se-SG), an
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oxidized form of glutathione (GSSG) (hazardous compound). GSSG is detoxified to the reduced
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form (GSH) by glutathione reductase in the vacuole. Glutathionyselenol (GS-Se-H) and hydrogen selenide (H2 Se) are formed because of further spontaneous metabolism of GS-Se-SG
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couple with GSSG. Via passive transport, H2 Se is secreted (through the vacuolar membrane) out into the cytoplasm of the cell. The activities of superoxide dismutase further synthesized
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glutathionylselenol into products (elemental Se and glutathione) (Tarze et al., 2007; Lazard et al. 2011; Mapelli, Hillestrøm, Kapolna, Larsen, & Olsson, 2011; Kieliszek et al., 2015). H2 Se is
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highly considered as a potent reducing agent along with glutathionyselenol (GS-Se-H), glutathione, and cysteine molecules (Kieliszek et al., 2015). H2 Se act as a precursor in the formation of organic compounds, including selenoamino acids where it is bound to O-acetyl homoserine (O-Ac-HSer) involving homocysteine synthase. Selenohomocysteine (SeHCys) and acetic acids are then synthesized (homoselenocysteine biosynthesis). The reaction is further branched into selenocystathionine or selenomethionine (SeMet) synthesis. SeHCys is bound to serine, which is then catalyzed by cystathionine βsynthase, forming selenocystathionine and water as a by-product. SeMet is substituted with formethionine because it easily acylates Met-tRNA which leads to the accumulation of
ACCEPTED MANUSCRIPT methionine-requiring proteins (The reaction is catalyzed by homocysteine methyltransferase). Another possible route is via trans-sulfuration mechanism, which is catalyzed by γ-lyase into selenocysteine (SeCys) and H2 Se (Pedrero & Madrid, 2009; Mapelli et al. 2011; Kitajima & Chiba 2013). The biosynthesis of SeCys in yeast cells is well elucidated in the following literature (Xu et al., 2007; Squires & Berry, 2008; Allmang, Wurth, & Krol, 2009; Turanov et
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al., 2011). Na2 SeO 3 reacts with reducing sugars in wort, forming the red elemental Se, which is toxic.
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(Mapelli et al., 2011). This toxic compound could end up in the Se-enriched beer. The ability of
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yeast to bind sulfur onto its cell wall is of interest to brewers. Sulfur act as a precursor for the formation of unpleasant flavors in some beers, which leads to loss of revenue since such
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products are poorly patronized.
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Rhodotorula mucilaginosa-13B could be used in cleaning water with a high Se dose due to its ability to biotransform and accumulate Se on its structures (Ruocco, Chan, Hanson, & Church,
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2014). S. uvarum can biotransform inorganic Se to organic form when pitched in nutrient-rich wort. This theory was confirmed in the work of Marinescu, Stoicescu, & Teodorof (2011) where
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they reported that Se became organically bound to the yeast when brewery wastewater was used as the medium for cultivation.
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It was reported by Sánchez-Martínez et al. (2012) that increasing the Se concentration in wort resulted in an increase in the percentage of Se in beer. However, above 10 µg/mL, the quantity remained the same. This result also depends on the kind of yeast (top or bottom fermenting) used. Thus, at a certain threshold, the yeast could no longer tolerate the toxicity of the incorporated Se. Therefore, this threshold should be deciphered so that brewers can manage and produce a better beer. This could also avoid brewing beer contaminated with toxic Se. Zare and
coworkers
(2017) reported that among the numerous yeasts screened for
biotransformation of inorganic Se to it bioavailable form, only S5, and S18 were found to yield good SeMet content (above 2500 ppm organic Se). Nevertheless, S18 isolate showed the
ACCEPTED MANUSCRIPT maximum biomass production and SeMet accumulation (2655 ppm) and (3.73 g/L) when exposed to Se at 25 mg/L concentrations. Pérez-Corona et al. (2011) reported the biotransformation of inorganic Se by S. cerevisiae and S. bayanus when they produced white wine under laboratory-scale. It was observed that the viability of cells in the must (must and inorganic Se) were lower compared to the control (YEPD
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medium) experiment, though this did not affect the entire fermentation process. The addition of 200–500 µg of SeO3 2- in the must resulted in the production of Se-enriched wine (55–60%, Table
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1). This percentage starts to decline (by approximately 30%) when the amount of SeO 3 2-
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increases, hence the maximum quantity of SeO 3 2- to incorporate in any beverage should be in the range between 200–500 µg. A scale-up experiment is warranted in order to validate the findings S. cerevisiae has higher biotransformation efficiency than the S. bayanus. Nevertheless,
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above.
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screening for Se toxicity level (tolerance level) of various yeast is a good initiative in selecting efficient strains for producing Se-enriched beverages with good sensorial properties and high
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acceptance levels. Since beer is one of the most consumed beverages in the world, Se beer can be considered an adequate source of Se for people deficient in Se.
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2.2. Production of selenized kvass
Kvass was invented by the traditional Slavs and is a favorite low alcoholic beverage in the
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Eastern European countries (especially in Russia and the Ukraine) and the former Soviet republics. It is produced from either rye/wheat bread (dried). The hue varies (color of bread to dark black) due to the utilization of different raw materials in the production process. By the Russian standard, the beverage is categorized as a non-alcoholic drink (0.5–1.0%). The demand for different flavors in kvass has caused producers to start adding fruits and other flavor enhancing substances (Hornsey, 2003; Sandor, 2003; Volhina, 2011; GOST, 2005). The entire production process of selenized kvass is summarized in figure 2. Rye grains were soaked and germinated in solutions of Na2 SeO 3 at room temperature for 4 days. The germinated
ACCEPTED MANUSCRIPT grains were then kilned in an oven to halt the enzyme activities. The enzymatic activities in the grains are thought to biotransform the absorbed SeO 3 2-. The malted rye grains were dried and weighed before crushing in a roller mill. The milled grains were then sieved through a screen with 1 mm openings (determined by weighing after milling) for better extraction during mashing.
Milled grains (500 g) was mixed with potable water and
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heated at 58 °C for 20 minutes. The ratio of the malt and water was 1:2.5. The temperature was increased to 64 °C and 72 °C with step times of 60 and 120 mins respectively. The temperature
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was finally increased to 78 °C for 10 mins. Filtration was carried with the aid of a cheesecloth o
Brix for the control and the Se-
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and 3.51 L (80 °C) of water was used for the sparging. The enrich wort was 11.7 and 12.4 respectively.
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Dry Se-biofortified pak-choi cabbage (3.69 g) (the concentration of the Se was 250 μg/kg) was
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ground to the size of 05.2 mm, water was then added at a ratio of 1:12. Enzyme preparations were made to enhance the extraction.
Distizim-protacid extra (manufactured by Erbsloh,
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Germany) and Viscostar (Novozymes manufacturer) had proteolytic and cellulolytic activities on the sample. The dose of enzymatic preparations was 1% per 1 kg of raw material. The
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temperature of the mixture (enzymes + sample) was raised to 55°C and held for 24 hours, the solution was filtered, and the Se concentration of the extract (filtrate) determined. The Se-
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enriched extract was poured into the wort. Fruit yeasts (Lesaffre) were reactivated before pitching for reparation of vital functions in stored yeasts, together with LAB Evitalia (1.25 g yeasts, 25 mg bacterial culture).
Fermentation lasted for 24 hours at 30 °C. Finally, the selenized kvass underwent pasteurization for 20 minutes at 80 °C (Antoņenko, Duma, Kreicbergs, & Kunkulberga, 2016; Muravyov, Barakova, Hamsters, & Panova, 2018a; Muravyov, Barakova, Hamsters, & Panova, 2018b; Muravyov, 2018).
It was allowed to cool before bottling followed by sensorial analysis
(selenized kvass). The Se-enriched extract could have been added to the green beverage before conditioning and not prior to fermentation. Quantification of Se in rye and the final products
ACCEPTED MANUSCRIPT were done according to Russian standard (GOST, 2008). Other techniques could be applied as well. The results revealed that selenized kvass produced from rye grains soaked and germinated in solutions of Na2 SeO 3 had the higher content of Se (51.1 μg/kg) when compared to the control experiment. However, the alcohol content (0.7%) was more than the control (0.5%), indicating that Se influenced the enzymatic activities in the malt. According to Antoņenko et al. (2016), the
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addition of Se significantly increased the activity of amylases. This might have increased the sugar content in wort, explaining the higher alcohol percentage recorded in the selenized kvass.
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3. Nutritional supplements (selenized-yeast (Se-yeast) and algae (Se-algae))
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Selenium-enriched supplements are ideal intervention to increase the Se intake for a large population, thus minimizing the risk of diseases and deficiencies associated with Se.
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The active groups of proteins, phospholipids, or polysaccharides of yeast cell wall acts as the
composed
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biological binder for Se (Kieliszek et al., 2015). According to the literature, the cell wall is of polysaccharides (85%) and
proteins (15%). Further examination of the
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polysaccharide revealed that it included glucose (80–90%), mannose (10–20%), and Nacetylglucosamine (1–2%) (Klis, Boorsma, & De Groot, 2006).
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The architecture of the cell wall is composed of inner layers of the carbohydrate polymer β-(1,3) glucan (30–45%) and the outer layers of mannoproteins (30–50% of the cell wall). Additionally,
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β-(1,6) glucan (8–18%) and chitin amount to 1-2% of the total composition of the cell wall (Klis et al., 2006; Saluk-Juszczak, Królewska, & Wachowicz, 2010; Levin & Moran 2011; Orlean, 2012).
Se-yeast is the product of the aerobic fermentation of various yeast species in a Se-enriched medium (Rayman, 2004). Yeast can bioaccumulate up to 3000 µg/g of organic Se (Schrauzer, 2006) which depends on the type of media, the conditions, and concentration of the media constituents. Inorganic Se is transformed into SeMet, which is the main selenoamino acid found in the enzymatic hydrolysis of yeast (Ponce de León et al., 2002; Rayman, 2004; Reyes et al., 2004;
ACCEPTED MANUSCRIPT Schrauzer, 2006). S. cerevisiae is one of the best organisms that can biotransform inorganic Se to the more bioactive SeMet form (Zare et al., 2017). Under stress conditions (high temperature, fluctuation in pH, etc.), yeast can convert inorganic Se into organic Se (SeMet) which can be accumulated on the cell wall. Increasing the scale of production of Se-enriched yeast is the last step in order to meet human
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demands for SeMet (Yin, Chen, Gu, & Han, 2009; Sanchez et al., 2012; Bierla, Szpunar, Yiannikouris, & Lobinski, 2012; Rajashree & Muthukumar, 2013b). However, green (clean)
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methods should be adhered to when fulfilling this goal in order to avoid cultivating yeast
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contaminated with heavy metals and other naturally occurring yeast strains. Bioaccumulation of SeMet (300 to 2200 ppm) was achieved with yeast cells using a synthetic medium. The yield was
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increased by supplementing the culture medium with Na2 SeO 3 (Suhajda et al., 2000; Antoneta,
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Marinescu, & Teodorof, 2011; Muthukumar & Rajashree, 2013; Zare et al., 2017). Many industries across the world have now mastered the large-scale production of Se-yeast. Its
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relatively low production cost makes it a promising and attractive venture for the food and pharmaceutical industries. A mathematical model was applied to study the effects of culture
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conditions (temperature, fermentation time, initial pH value, shaking speed, as well as time and concentration of Se, added to medium) on the bioaccumulation of Se in yeast. The total Se
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accumulation and organic Se formation ranges from 107.9 to 287.6 mg/kg and 93.27 to 269.05 mg/kg, respectively and the best conditions to obtain these results were: Se concentration, 25 μg/mL; addition time of Se source, 9 h; inoculums size, 30 g/L; shaking rate, 130 rpm; fermentation time, 48 h; temperature, 28 °C; and pH, 5.8 that have maximum Se incorporation (Esmaeili, Khosravi-Darani, Pourahmad, & Komeili, 2012). Se-yeast is the most studied Se-enriched product. However, Se species identification and quantification in this matrix are still a challenge (Pedrero & Madrid, 2009). Recent development has helped to solve this shortfall (Méndez, González, & Sanz-Medel, 2000; McSheehy, Pannier, Szpunar, Potin-Gautir, & Lobinski, 2002; Hinojosa-Reyes, Ruiz-Encinar, Marchante-Gayun,
ACCEPTED MANUSCRIPT Garcia-Alonso, & Sanz-Medel, 2006). Yeast Candida utilis ATCC 9950 was found to secrete Se in culture media which is triggered by enzymes associated with intracellular transport of Se. This leads to bioaccumulation of organic Se in the cell wall, which is a good source of dietary Se. Electrophoresis of the medium revealed protein fractions of yeasts obtained from media supplemented with Se, the presence of new protein fractions characterized by molecular weights
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of 60, 22, 18, and 15 kDa (Kieliszek et al., 2015). However, high accumulation of Se in S. cerevisiae yeast cells occurred in protein fractions above 75 kDa (Bryszewska, Pęczkowska,
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Rudzi´nski, Diowksz, & Ambroziak, 2000). The biosynthesis of selenoproteins in yeast cells is
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induced due to the presence of Se in the medium (Kieliszek et al., 2015), which are the bioaccessible form of Se. Se-yeast could be utilized in formulating different kind of foods i.e. ice
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cream, porridge, drinks, etc.
Se-enrich medium.
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According to Rayman (2004), Se-enriched yeast cream is the product after culturing yeast in a Pasteurization is then carried out, thereby killing the yeast, followed by
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spray drying. Dried Se-yeast can be used in formulating foods like ice creams and cereal basedcomplementary foods. Stabnikova et al. (2008) added a solution of sodium hydroselenite
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(NaHSeO 3 ) with a concentration of 100 µg Se/mL to the sterile media utilized for incubating S. cerevisiae. The growth (kinetics) and influence of Se on yeast biomass at different
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concentrations were studied. It was established that the growth was inhibited due to the addition of NaHSeO 3 to the medium, and resulted in the accumulation of elemental Se, which is not suitable for human consumption. The specific growth rates declined (0.146 to 0.073 L/h) when the Se concentration increased (from 2 to 12 µg/mL). The obtained Se-enriched yeast was used to bake Se-enriched bread. Low fermentative activities of yeast and longtime of the dough pellet to float up (rise power) were some of the problems faced by the authors. Nevertheless, they curb these problems by re-suspending (maturation) the Se-enriched yeast in the solution of NaCl, 9 g/L that significantly improved the baking properties. The 100 g of Se-enriched bread contained up to 50 µg Se in the form of SeMet.
ACCEPTED MANUSCRIPT According to Lazo-Vélez, Chávez-Santoscoy, & Serna-Saldivar (2015), the consumption of Seenriched bread could counteract deficiencies - especially in regions that have poor Se soils. However, several factors (i.e., the agronomic, milling, and processing, etc.) significantly affected Se concentration and bioavailability of the flour. Hart et al. (2011) utilized Se biofortified wheat grain cultivated by Broadley et al. (2010) in
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making bread flour for baking Se-enriched bread. A linear correlation was established between the dose of Na2 SeO 4 applied to the crop and the amount of Se in flour and bread indicating a
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minimal loss of Se during grain processing and bread production. SeMet was the predominant Se
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species in the products (flour and bread) obtained from wheat grain biofortified with Se fertilizers. However, SeCys, Se-methylselenocysteine, SeO 3 2-, and SeO 4 2- were other Se species
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detected in the Se-enriched flour and bread. Applying 10 g/ha of Na2 SeO 4 fertilizer raised the
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total Se in wholemeal and white bread by 185 and 155 µg/g, respectively. Supplementing dairy (n = 42) cattle feed with Se-yeast has been shown to increase the amount of
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milk, thus increasing cheese production. The milk Se concentration reached a plateau within a week after commencing Se supplementation. Se-yeast was more effective than inorganic Se
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(Malbe et al., 1995; Ortman & Pehrson, 1999). According to Dillehay et al. (2008), algae have been part of human civilization for thousands of
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years, based on archaeological evidence from 14,000 calendar years before the present (cal yr B.P.) in Chile. China and Indonesia ranked as the top producers of seaweeds and is estimated to value about US $6.7 billion. However, these algae are utilized as foods and food additives (Switzer, 1980; Jassby, 1988; Fournier, Adam, Massabuau, & Garnier-Laplace, 2005; Gantar & Svircev, 2008; Chacón-Lee & González-Mariño, 2010; FAO, 2015; FAO 2016). The demand of algal foods varies across the world due to culture, for instance, typical Japanese diets contain about 9.6 (2014) to 11.0 (2010) g macroalgae per day (MHLW, 2014). Globally the demand for macroalgal and microalgal foods is growing, and they are increasingly being consumed (Wells et al., 2017) or used in formulating functional foods due to its high nutritional value. The schematic
ACCEPTED MANUSCRIPT model of Se metabolism in the algal cell is depicted in figure 3. Detailed Se accumulation and metabolism in algae is reviewed in (Schiavon, Ertani, Parrasia, & Vecchiad, 2017). In aquatic ecosystems, microalgae act as a significant vector of Se accumulation from the watercolumn and partially transform it into organic Se before it is transferred by ingestion to higher organisms. Among the species of microalgae, Spirulina platensis is found to bioaccumulate Se at
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high concentration as 400 mg Se/g dry weight on their cell walls. Scenedesmus quadricauda is reported to also accumulate high Se in the form of Se-enriched microalgal biomass (Fan, Teh,
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Hinton, & Higashi, 2002; Douskova´ et al., 2007; Chen, Zheng, Wong & Yang, 2008; Doucha,
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Lıvansky, Kotrbacek, & Zachleder, 2009).
The European Space Agency, and the National Aeronautics and Space Administration (NASA)
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in the late 1980s and early 90s proposed Spirulina as one of the primary foods to be cultivated
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during long-term space missions due to its nutritional value (Tadros & MacElroy, 1988; Cornet & Dubertret, 1990). Growth media (conditions, i.e., pH, O2 , etc.) can be manipulated to
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significantly increase the nutritional value of algae (Tadros & MacElroy, 1988). Therefore, adding inorganic Se into the growth media could produce Se-enriched algae, which has a great
populations.
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potential of increasing the intake of bioavailable form (organic) of Se by selenium-deficient
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Unicellular marine algae (Dunaliella primolecta, Porphyridium cruentum, and Chlorella vulgaris) have been reported to be excellent accumulators of Se (Bottino et al., 1984; Sun, Zhong, Huang, & Yang, 2014). A significant increase in the cell growth rate and organic Se level of C. vulgaris was reported when exposed to different concentrations of Na2SeO3. Se positively promoted C. vulgaris growth at lower concentrations (≤75 mg L-1), acting as an antioxidant through the inhibition of lipid peroxidation (LPO) and intracellular reactive oxygen species (ROS). The Se-Chlorella could be utilized in designing antioxidative functional foods human health (Sun et al., 2014).
ACCEPTED MANUSCRIPT Spirulina is widely used around the world, because of its rich source of vitamin B12 , in addition to vitamins B1 and B2 . It is estimated to provide one-half of the adult daily requirements of vitamin A (B-carotene) (Tadros & MacElroy, 1988). Arsenyeva & Skripleva (2014) designed yogurt enriched with the supplement «Selenium Algae Plus». A highly bioavailable organic form (SeMet, SeCys) of Se was found to be present in this
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supplement and has a digestibility of 95-98%. The assimilation of Se was found to be inhibited in the presence of sucrose. Therefore, Jerusalem artichoke syrup and stevioside were substituted
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as alternative sweeteners to curb this issue. Nevertheless, these sweeteners can reduce the level
for people who have diabetes and other ailments.
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of sugar in the blood further. The Se-enriched yogurt is not only for mass consumption but also
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The geographical locations, seasons and culture practices (aquaculture) are obstacles in
achieving specific objectives.
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understanding the Se and other nutritional constituents of algae, which impede their utilization in
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POLG mutator mice were used as model organisms during feeding trials to establish the effects of dietary Se status on cardiac gene expression. Diet supplementation significantly (P <0.05, FC
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>1.2) suppressed the expression of age-induced genes that are functionally related to cardiomyocyte apoptosis, hypertrophy, and fibrosis, probably by regulating the transcriptional
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activity mediated by Foxo transcription factors (Xiao et al., 2014). Food processing and storage could potentially alter the nutritive value of algal-formulated foods. Supplementing different forms of Se (Na2 SeO 3 , Se-enriched yeast, lactate protein Se complex, and Se-proteinate) in the feed of pregnant goats revealed elevated Se concentration in the blood of treated animals when compared to the control group. Additionally, kids (about 60% of Se) from treated mothers had higher Se levels in the blood than kids from the control group (Pavlata, Mišurová, Pechová, & Dvořák, 2012). Chlorella sorokiniana is another alga species that can bioaccumulate Se for designing functional food. A Na2 SeO 4 -enriched medium was applied in optimizing growth conditions of C.
ACCEPTED MANUSCRIPT sorokiniana. It was revealed that Se concentration of 100 μg/mL in the medium was the maximum concentration at which growth was maximal. Increasing the concentration above 100 μg/mL caused the collapse (death) of the colonies. Moreover, SeO 3 2- in the medium was biotransformed into mainly SeCys, selenomethyl-selenocysteine (SeMeSeCys), and SeMet, which was confirmed by metallomics analytical approach coupled with HPLC-ICP-MS (Gómez-
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Jacinto, García-Barrera, Garbayo, Vílchez, & Gómez-Ariza, 2012). Microcystins (MCs) in Spirulina are toxic and pose a potential threat to consumers. Roy-
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Lachapelle, Solliec, Bouchard, & Sauvé (2017) detected cyanotoxins in 18 brands of
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Cyanobacteria dietary supplements containing Spirulina and Aphanizomenon flos-aquae. The risks of consuming these products were also assessed. It was discovered that 3 brands out of the
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14 Spirulina products contain MCs ranging between 0.25 and 0.84 µg/g, whereas, 0.8 and 8.2
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µg/g was associated with the A. flos-aquae-based brands. The sum of Spirulina and A. flosaquae-based products MCs varied between 0.01 and 0.63 µg/g and 0.52 and 5.8 µg/g,
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respectively. It was concluded that the quantity of MCs in these brands exceeded the recommended tolerable daily intake (TDI) for adults specified by the World Health Organization
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(exceeding TDI up to 683% for total MCs) (WHO, 2011; Roy-Lachapelle et al., 2017). The amino acid phenylalanine in spirulina causes brain damage - especially to people suffering from
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phenylketonuria (Robb-Nicholson, 2006; Habib, Parvin, Huntington, & Hasan, 2008; Heussner, Mazija, Fastner, & Dietrich, 2012). These toxins could be minimized if organic fertilizers are utilized in cultivating these algae. Chemicals or other additives should be avoided during growing and handling (processing) of algae. There are reports of massive metal accumulation in spirulina supplements and were confirmed in a recent study (approximately 5.1 ppm of the lead (Pb) was detected) (Siva-Kiran, Madhu, & Satyanarayana, 2015). Constructing the cultivation site (pond) far away from industrial settings where the air is less polluted with the residue of heavy metals coupled with using fresh water could be the ultimate remedy for this menace. Nevertheless, using a closed photobioreactor where the environment (internal and external) is
ACCEPTED MANUSCRIPT easily controlled could be an alternative approach. The device could aid in controlling cyanobacterial bloom, therefore minimizing the risk of MCs in the product. The exact doses of MCs are still an open issue for debate, which should be solved in order to prevent possible health risks. Therefore, enacting national and international legislation is necessary to preserve aquatic environments as well as human health (Drobac et al., 2013).
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Se-alga could be useful in designing new functional foods that could exert positive health benefits to consumers. This could be in the form of anti-cancer effects, increase in energy,
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boosting the immune systems, antioxidant proprieties, etc. that will help save huge sums that
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would have ended up in seeking medical care. It could also act as a precursor for selenoprotein synthesis.
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However, in reality, the widespread use of selenized supplements in formulating functional foods
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has a specific limitation. Firstly, the addition of supplements could alter the organoleptic properties of the product. Secondly, some consumers might be allergic (react) to some of these
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products.
According to Rayman (2004), the initial production of selenized supplements was not meant for
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only human consumption. However, the increase in demand for these products across the world following the approval from the US Food and Drug Administration has shifted perception on
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these supplements. The leading manufacturers of these supplements are Pharma Nord, Vejle, Denmark, Alltech, Lexington, KY, USA, Lallemand, Montreal, Canada, etc. Manufacturers are cautioned to follow international standards in order to avoid contaminations, i.e. microbial or heavy metals.
The tolerable upper intake level (UL) of Se for adults in the US is 400 µg (5.1 µmol)/day based on selenosis as the adverse effect. UL for infants of the age 0–6 and 7–12 months are 45 µg (0.57 µmol)/day and 60 µg (0.76 µmol)/day of selenium, respectively. Children of the age 1–3, 4–8, 9– 13 years has UL of 90 µg (1.1 µmol)/day, 150 µg (1.9 µmol)/day, and 280 µg (3.6 µmol)/day of selenium, respectively (Food and Nutrition Board Institute of Medicine, 2000).
ACCEPTED MANUSCRIPT The European Commission health and consumer protection directorate have derived the UL of 300 µg Se/day for adults, pregnant and lactating women based on no-observed-adverse-effect level (NOAEL) of 850 µg/day for clinical selenosis in the study on 349 subjects of Yang et al. (1989b) and Yang & Zhou (1994). In addition, UL of 250, 200, 130, 90, 60 μg selenium/day was derived for Europeans of the age 15-17, 11-14, 7-10, 4-6 and 1-3 years respectively (European
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Commission Health & Consumer Protection Directorate, 2000; European Food Safety Authority, 2006). Some European Union countries have set the mean daily intakes of Se for non-vegetarian
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adults as: United Kingdom 63 μg/day, France 29-43 μg/day, Sweden 24-35 μg/day, Belgium 28-
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61 μg/day, Denmark 41-57 μg/day, Finland 100-110 μg/day, The Netherlands 40-54 μg/day, Norway 28-89 μg/day, and Spain 79 μg/day (Alexander & Meltzer, 1995; van Dokkum, 1995;
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Johansson et al., 1997). The Se fertilization programme in Finland appeared to have successfully
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raised Se status in the population (100-110 μg Se/day). However, in the case of Norway, this could be due to the continuous importation of biofortified wheat (European Commission Health
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& Consumer Protection Directorate, 2000; Stoffaneller, & Morse, 2015). The recommended UL by the Americans (400 µg) is higher when compared to the EU value (of 300 µg Se/day).
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3.1. Preparation of selenized Tom-brown porridge flour Tom-brown flour is a powder made by grinding roasted raw grains and is used for making
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different foods. This flour is the main ingredient for porridge which nursing mothers used in weaning their babies in Africa. Nursing mothers could also use this porridge in supplementing the breast milk for their babies since it expensive buying products like milk (NAN milk), cerelac, etc., which are formulated in other parts of the world. Figure 4 shows the entire process of producing the Tom-brown porridge flour. 1. After harvesting and drying of cereals (yellow corn and guinea corn). Screening is followed to remove stones and other foreign particles. Further cleaning such as winnowing is carried out. The cereal is then roasted in a frying pan on a fire until it turns golden brown. It is then removed and allowed to cool.
ACCEPTED MANUSCRIPT 2. Depending on the location, legumes like groundnuts, soybeans, etc. could be added in the mixture. Similar treatments as explained above are replicated here. 3. The roasted cereals and the legumes are mixed and milled into flour using either an electric or a diesel-milling machine. Some producers do pound the grains in mortar and pestle to separate the chaff. However, they are criticized of eliminating vitals nutrients
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that might have attached to these chaffs. 4. Pasteurized dried selenized supplements (yeast and algae) are then mixed with the milled
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flour (Selenized enriched Tom-brown).
4. Extraction and quantification of organic Se.
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5. The selenized enriched Tom-brown flour is now ready for consumption.
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Several researchers have outlined methods for quantifying Se in various matrixes (Uden,
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Boakye, Kahakachchi, & Tyson, 2004; Kaplan, Gil, & Cerutti, 2006; Harwood & Su, 1997; Goenaga-Infante et al., 2008; Pedrero, & Madrid, 2009; Soruraddin, Heydari, Puladvand, &
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Zahedi, 2011; Chen et al., 2011; Nam, & Basu, 2011; Duan, He, & Hu, 2012; Fagan et al., 2015; Jagtap & Maher, 2016; Tie et al., 2016). The chemical form of Se in sample, matrix (i.e., plant,
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fish, meat, yeast, egg, human serum, blood, etc.), the instrumentation availability, experience of the researcher are factors to consider when selecting methods for sample treatment for the
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isolation and identification of the Se species (Pedrero & Madrid, 2009). According to Peachey, McCarthy,
& Goenaga-Infante, (2008) selecting extraction methods, which exhibit high
extraction efficiency while preserving the integrity of Se species is essential for the accurate measurement of Se species. Extraction of Se species using aqueous solutions (water, water– MeOH, and neutral buffers), acid–alkali hydrolysis (HCl, methanesulfonic acid, tetra-methylammonium hydroxide (TMAH)), and enzymatic hydrolysis (proteinase K, subtilisin, pepsin and protease, or mixture of enzymes protease and lipase) are the methods available currently. Acidic extraction is characterized with low SeMet recoveries, in some instance complete loss of SeCys (~100%) with extracts containing elementary Se0 (Huerta, Sanchez, & Sanz-Medel, 2004; Yang,
ACCEPTED MANUSCRIPT Sturgeon,
McSheehy,
& Mester,
(Selenomethyl-selenocysteine,
2004; Jagtap
& Maher,
γ-glutamylselenomethyl-selenocysteine,
2016). Selenoamino acids etc.)
are
water-soluble
hence, are aqueously extracted with hot water with excellent recovery. However, excellent recovery is only achieved if Se species are not incorporated into proteins (Pedrero & Madrid, 2009). Therefore, coupling aqueous with enzymatic treatment could improve the extraction
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efficiency. Enzymes can interact, creating pores on proteins housing Se species, thereby resulting in leaching. Organic Se was successfully extracted from selenized Yeast (McSheehy, Szpunar,
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Haldys, & Tortajada, 2002; Encinar et al., 2003), plants (Zhang & Frankenberger, 2001) and
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mushrooms (Stefanka, Ipolyi, Dernovics, & Fodor, 2001) using an aqueous method with good recoveries. Previous findings (Moreno, Quijano, Gutiérrez, Pérez-Conde, & Cámara, 2004;
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Huerta, Sanchez, & Sanz-Medel, 2004; Dumont, Vanhaecke, & Cornelis, 2006) have suggested
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that the incorporation of sodium dodecyl sulfate (SDS) increased/improved the extraction efficiency up to 50%. SDS interact with proteins thereby rendering them soluble in water hence
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increasing the release of selenoproteins. Chen et al. (2011) modified the method previously described by Dumont et al. (2005) to isolate Se species from both yeast and clover. High purity
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deionized water (5mL) was added to a 25 mg sample in a centrifuge tube, followed by a hot water bath at 50 ◦C for 24 h. The extract was then centrifuged and filtrated, and the filtrations
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were made up to 10mL with deionized water. The enzymatic hydrolysis is the preferred method of many researchers because it presents several merits compared to the other methods. Enzymatic hydrolysis is selective and specific because enzymes act only on specific chemical bonds and can therefore distinguish between fractions of elements associated with the different components of the sample matrix. Furthermore, elemental losses by volatilization are minimized because enzymes perform well at moderate pH and temperature.
Reagents are not required to neutralize the excess of acid or
alkali in acidic extracts before chromatography analysis hence reducing any risk
of
ACCEPTED MANUSCRIPT contamination (Bermejo-Barrera, Fernandez-Nocelo, Moreda-Pineiro, & Bermejo-Barrera, 1999; Jagtap & Maher, 2016). The Se yeast sample could accurately be measured (0.04 g) into microcentrifuge tubes. Protease enzyme solution (13.33 mg Protease XIV in 0.5 mL of Tris buffer pH 7.5) is then added and vortex mixed for 2 min. Samples were ultrasonicated for 25 s at 80 % amplitude, a power output
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of 30 W for 15 minutes. Ice cubes is used to cool the sample from the heat generated due to the ultrasonication. Before commencing the sonication, 250 μL enzyme solution is used to rinse the
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probe. Extracted samples is then centrifuged at 14,000 rpm for 3 min. Supernatants are then
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transferred to 15 mL centrifuge tubes. Pellets washed with water (0.9 mL) and vortex-mixed completely in solution. This step is repeated thrice. Volumes are adjusted to 15 mL using water
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and mixed thoroughly before removing an aliquot (2 mL) for filtration (0.25 μm) and dilution
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before analysis (Fagan et al., 2015). Residual yeast, white wine, and beer were subjected to enzymatic hydrolysis before quantification and identification of selenium species (Pérez-Corona
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et al., 2011; Sánchez-Martínez et al., 2012). The extracts are then injected into high-performance liquid chromatography with inductively coupled plasma mass spectrometry (HPLC-ICP-MS)
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using a sample injection valve fitted with a 100 µL loop. Details of the separation columns and operating conditions are reported in Pérez-Corona et al. (2011) and Sánchez-Martínez et al.
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(2012). Comparing the retention time and spiking experiments Se species are identified. Se species (Se(VI) and SeMet from impurities in the protease hydrolysis is used as blank for the analysis. High-resolution Time-of-flight Mass Spectrometry could also be used in quantifying organic Se (Krittaphol et al., 2008). Good Se recoveries by enzymatic hydrolysis of yeast, fish and plant CRMs and miscellaneous have been reported. Approximately 96% and 93% of total Se in Antarctic krill and cod muscle (CRM BCR 422) were documented, respectively (Siwek, Galunsky, & Niemeyer, 2005; Kapolna, Gergely, Dernovics, Illes, & Fodor, 2007). Hydride generation–inductively coupled plasma optical emission spectrometry (HG-ICPOES) was proposed as an effective method of
ACCEPTED MANUSCRIPT determining total Se in the sample. This system allows the determination of Se in high copper dose samples. High precision and accuracy were obtained after analyzing reference samples (Kaplan, Gil, & Cerutti, 2006). The outlined approaches provide a secure, rapid, reproducible, and cost-effective platform for computing organic Se. However, acidic hydrolysis of Se-enriched foods was found to be the most effective technique of measuring total Se. Therefore, coupling
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two pretreatment methods are recommended for effective extraction of Se species from samples. 5. Se bioavailability
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Ruby et al. (1999), defined bioavailability as the fraction of a substance released from food
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matrix that is absorbed via the intestinal barrier (bioavailable fraction), transported into the bloodstream or an organ, and is available to promote its activities in the exposed organism (Ruby
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et al., 1999). However, the non-accessible fraction is not absorbed, thus is directly excreted
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(Moreda-Pineiro, Moreda-Pineiro, & Bermejo-Barrera, 2017). The fraction of a nutrient or substance in food that is theoretically soluble from its matrix and can quickly enter the
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bloodstream or affect the gastrointestinal (GI) tract is termed bioaccessibility (Ruby et al., 1999; Oomen et al., 2002; Stahl et al., 2002; Moreda-Pineiro et al., 2017). The physiological state of
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the consumer (i.e., age, health status, gender, etc.,), GI conditions, the form of Se in food (Ziegler et al., 1978), and geographical location are factors affecting the bioavailability of any
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food.
With the aid of isotopically or non-isotopically labeled Se (Miller et al., 1981; Ruby et al., 1999; Fernandez-Garcıa et al., 2009) several stimulated studies have been carried out to assess the bioavailability of Se in natural products or Se-enriched, raw and processed products. Recently Moreda-Pineiro et al. (2017) published a good review on Se bioavailability in various food products. In vivo, human tests show a significant increase in the Se levels in plasma or whole blood after supplementation
with
Se-fortified
wheat
and
process-fortified
Se
biscuits.
Plasma
Se
concentrations with Se-fortified wheat was more than (from 122 to 194 µg/L) with process-
ACCEPTED MANUSCRIPT fortified Se biscuits (from 122 to 140 µg/L). This could be as a result of different species of Se available in the products (i.e., SeMet and SeOMet in Se-fortified wheat and process-fortified Se biscuits, respectively) (Thompson et al., 1985; Mutanen, 1986; Van der Torre et al., 1991; Djujic et al., 2000; Kirby et al., 2008). A similar study was carried out on rats using tuna and Sefortified wheat. There was a significant decrease in glutathione peroxidase (GSHPx) activity
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with groups treated with Se-fortified wheat compared with the tuna treated groups, suggesting that Se in wheat was more available than that in tuna (Alexander, Whanger, & Miller, 1986).
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Yoshida and colleague (1999) quantified tissue Se accumulation and GSHPx activity of 4-week-
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old male Wistar rats fed with high-Se yeast (SeY) and Na2 SeO 3 . Strong correlations were established between the supplementary Se levels and the tissue Se contents or GSHPX activities,
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indicating that that Se in SeY is more bioavailable than Na2 SeO 3 . Therefore, SeY should be used
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in formulating Se functional food.
Selenium-deficient rats showed dose-dependent increases in GSHPx activities in blood and in
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thioredoxin reductase activity in the liver after feeding them with yellow peas and oats harvested from the high-Se soil of South Dakota, the United States. It was concluded that Se from yellow
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peas and oats was highly bioavailable (Yan & Johnson, 2011). In a similar study, soy protein isolates (from soybean) was more Se bioavailable relative to tofu (Yan, Reeves, & Johnson,
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2010).
Bioavailability of Se (organic Se species such as MeSeCys, SeCys2, and SeMet) in Allium was found to be very high after an in vitro gastric and gastrointestinal digestion (proteolysis) than in Chive. Speciation studies revealed that these Se fortified Allium was not safe for human consumption because Se isolated was in the form of selenite. However, 90% of the total Se content of both samples became bioaccessible (Kapolna & Fodor, 2007). This finding was in agreement with previous studies (de Leon, Sutton, Caruso, & Uden, 2000; Kotrebai, Birringer, Tyson, Block, & Uden, 2000; Shah, Kannamkumarath, Wuilloud, Wuilloud, Caruso, 2004; Wrobel et al. 2004).
ACCEPTED MANUSCRIPT Selenized yeast and yeast-based nutritional supplements have been extensively assessed using in vitro approaches. Supplementation of the human diet with yeast and selenized tablets revealed that Se levels in plasma and whole blood increased significantly, indicating high Se availability in yeast (Alfihan, Aro, Arvilommi, & Huttunen, 1991; Xia, Zhao, Zhao, Zhu, & Whanger, 1992; Thompson, Robinson, Butler, & Whanger, 1993; Alfthan et al., 2000; Larsen et al., 2004; Bugel
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et al., 2008; Bost & Blouin, 2009; Mahmoud, 2012). In vivo animal (pig, ewe, rat, catfish, yellowtail kingfish, trout, and boiler chicken) studies showed that Se in yeast was highly
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bioavailable. Moreover, Se in yeast was reported to be almost twice as bioavailable as Se from
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Na2 SeO 3 and Na2 SeO 4 . High bioaccessible percentage (55–89%) was reported in the case of total Se in yeast, whereas low bioaccessible percentages (26–41%) were attributed to SeMet in yeast
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and yeast-enriched supplements (Smith & Picciano, 1987; Wang & Lovell, 1997; Yoshida et al.,
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1999; Dumont, Vanhaecke, & Cornelis, 2004; Hinojosa-Reyes, Ruiz-Encinar, Marchante-Gayon, Garcıa- Alonso, & Sanz-Medel, 2006a; Hinojosa-Reyes, Marchante-Gayon, Garcıa-Alonso, &
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Sanz-Medel, 2006b; Mateo, Spallholz, Elder, Yoon, & Kim, 2007; Wang & Xu, 2008; Rider et al., 2009; Rider, Davies, Jha, Clough, & Sweetman, 2010; Jang et al., 2010; Hall et al., 2012;
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Briens, Mercier, Rouffineau, Vacchina, & Geraert, 2013; Le & Fotedar, 2014). Hence, selenized yeast could be exploited to produce Se fortified functional food, which can serve as a dietary
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source of Se intake.
Food processing (i.e., cooking, frying, grilling, milling, fermentation, etc.) could affect the bioavailability of Se (Moreda-Piñeiro et al., 2017). Cooked tuna had slightly higher Se bioaccessible than in the uncooked sample. The heat from cooking might have affected the efficiency of digestive enzymes, hence improving protein digestibility, which facilitated Se release to enhance the bioaccessibility (Cabanero et al., 2004). In vivo, Se-deficient male albino rats showed higher bioavailability of Se in raw salmon than cured salmon. Curing salmon might have altered some enzymes, which could have inhibit the release of Se. However, Se from raw and cured salmon tended to be more bioavailable compared with Na 2 SeO 3 . The induction of
ACCEPTED MANUSCRIPT plasma GSHpx activity and Se accumulation in the femur and muscle goes to support the results (Ørnsrud & Lorentzen, 2002). Shi & Spallholz (1994) observed a significant decrease in liver GSHPx activity when Fischer 344 rats were fed a Se-deficient diet than the control rats. Upon supplementing the diets with 0.10 mg/kg Se as Na2 SeO 3 , Na2 SeO 4 , raw or cooked ground beef, liver GSHPx activity began to
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increase on treated rats than on the control rats. The percentage of liver GSHPx activity recovery were as follows: cooked ground beef (139%, p < 0.05), raw beef (127%, p < 0.05), Na2 SeO 4
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(117%, p < 0.05), and Na2 SeO 3 (98%, p > 0.05) indicating the bioavailability of Se from cooked
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ground beef and raw beef were greater than Na2 SeO 3 or Na2 SeO 4 . The cooking might have activated certain enzymes to hydrolyze all the peptide bonds in proteins, magnifying Se release
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thus increase its bioaccessibility. Milling has been shown to decrease the amount of Se in cereal
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product. The losses appear not to be excessive since Se is evenly distributed throughout the wheat kernel. Nevertheless, the Se that was lost to the consumer due to milling appeared in the
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by-products used as animal feeds, therefore some of this Se would eventually be in the meat and ultimately utilized by the consumer (Ferritti & Levander, 1974). According to Pederson et al.
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(1991) and Fairwetaher-Tait (1997), the presence of arsenic, vitamin C and guar gum (guaran) in the diet inhibit the uptake and assimilation of Se making it non-bioaccessible to the consumer.
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6. Mechanistic understanding of Se toxicity and health concern It is scientifically proven that organic Se (Se-yeast) is less (not toxic) than inorganic Se. Humans and animals (i.e., farm animals and poultry) are susceptible to excess or insufficient Se in the diet. High intake of organic and inorganic forms of Se have similar effects (Institute of Medicine, Food and Nutrition Board, 2000). The toxicity of Se depends on many factors, and it is difficult to make categorical pronouncements about the quantities of Se involved. The quantity and chemical forms of Se consumed coupled with other elements i.e. arsenic mercury, cadmium, etc. and dietary constituents could determine the class under which the toxicity (acute and chronic toxicities) is categorized (Diplock & Hoekstra, 1976; Farzaneh, 2015).
ACCEPTED MANUSCRIPT Seafood and meats (organ) are rich sources of Se in food. Other sources include muscle meats, grains, and dairy products (Table 2). However, care should be taken when choosing these products since some contain an excess amount of Se. For example, Brazil nuts contain around 68–91 mcg per nut and could lead to Se toxicity (poisoning) if consumed on a regular basis. In a study, when 201 people consumed a liquid dietary supplement containing 200 times Se than the
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recommended level, they experienced severe adverse reactions. i.e., vomiting etc. (Sunde, 2006; Sunde, 2010; MacFarquhar et al., 2010; Sunde, 2012).
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Saha, Fayiga, Hancock, & Sonon, (2016) discovered that the minimum lethal dose of Se in the
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form of Na2 SeO 3 varied from 1.5 to 8.0 mg/kg body weight in farm animals. Acute toxicity studies showed the LD50 for Se-yeast was 37·3 mg/kg compared with 12·7 mg/kg for Na2 SeO 3
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(toxic) (Figure 5). This evidence proved that Se-yeast is considerably less toxic than Na2 SeO 3
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(Vinson & Bose, 1987; Spallholz & Raftery, 1987).
When Se is consumed, be it inorganic (selenite) or organic (selenomethionine, selenized-yeast
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cells, etc.), it is metabolized by reduced glutathione (GSH) to yield seleno-diglutathione, which is then transformed into seleno persulfide (GSSeH). Under anaerobic conditions, GSSeH could
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enzymatically convert into volatile hydrogen selenide (H2 Se) or decays spontaneously to Se and GSH. Na2 SeO 4 is assimilated by active transport in the intestinal brush border whereas via
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passive diffusion, Na2 SeO 3 is absorbed. H2 Se enters the host cell with efficiency and exacerbates oxidation of GSH (Ganther, 1971; Hsieh & Ganther, 1975; Tarze et al., 2006). According to Shelly (2009), GSH is a ubiquitous intracellular protein with diverse functions that include detoxification,
antioxidant defense,
maintenance of thiol levels,
and modulation of cell
proliferation, etc. Se can combine with GSH and other thiols to form glutathioselenol, which could further reduce GSH into H2 Se. ROS such as superoxide ions (O 2 •), hydrogen peroxide (H2 O2 ), and hydroxyl radicals (OH•), are likely to be generated from the reaction between H2 Se and molecular cell oxygen. The reaction generates oxygen for the continuous oxygenation of glutathione resulting in the depletion of intracellular GSH, S-adenosylmethionine, which inhibits
ACCEPTED MANUSCRIPT protein synthesis, resulting in the complete substitution of sulfur (trans-sulfuration pathway) by Se in cellular metabolism. This leads to the loss of enzymatic activity in some sulfur-containing proteins due to the replacement of sulfur in sulfhydryl groups or thiols (vital for the formation of a disulfide bond) with Se. The consequence is genotoxic/cytotoxic effects due to oxidative stress on the cells (Vernie et al., 1974; Stadtman, 1974; Hoffman, 1977; Anjaria & Madhvanath, 1988;
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Seko, Saito, Kitahara & Imura, 1989; Stadtman, 1990; Yan & Spallholz, 1993; Spallholz, 1994; Seko & Imura, 1997; Pinson, Sagot & Daignan-Fornier, 2000; Tarze et al., 2006).
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Toxicity may also arise due to the reactions of H2 Se with metal-containing proteins, and this was
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confirmed by in vitro experiments. Recent studies showed that the iron atom in lipoxygenases of human monoclonal B-lymphocytes was sensitive to H2 Se. It was also reported that H2 Se
consequently affecting the human hepatoma cells (Eghbal,
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altering the redox reaction,
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exacerbates the inhibition of heme-containing enzymes, which belong to the respiratory chain,
Pennefather & O’Brien, 2004; Bjornstedt et al., 1996). These reactions could lead to dizziness,
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unconsciousness, etc. Pancreatic ribonuclease A was almost completely inactivated when treated with selenious acid (Ganther & Corcoran, 1969). Se toxicity could result in cell damages of
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genetic information (genotoxicity), which causes mutations. The availability of excess ROS in the cell reacts with both deoxyribose and bases in DNA, causing base lesions and strand breaks.
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Nevertheless, ROS can significantly oxidize DNA, thus inhibiting DNA repair, gene regulation (down-regulated) and threatening gene stability (Ramana, Boldogh, Izumi, & Mitra, 1998; Kelly et al., 2008; Wei et al., 2001; Zhou et al., 2003). In-vivo and in-vitro studies have confirmed that liver glutathione (thiols) producing organ is susceptible to Se toxicity due to its involvement in the rapid catalytic reaction of Se compounds. Thiol-containing enzymes like methionineadensyltransferase,
succinate-dehydrogenase,
NADP+-isocitrate-dehydrogenase,
and
lactate-
dehydrogenase were altered via Se toxicity (Nebbia et al., 1990; Mezes & Balogh, 2009). According to the literature, the minimum toxic dose of Na2 SeO 3 can cause clinical changes in birds, mg/kg body weight: ducklings – 9.4; broilers – 1.7; laying hens – 33.4; indices – 0.9. The
ACCEPTED MANUSCRIPT oral LD50 Se for young turkeys, ducks, chickens are 13.5, 64.0, and 9.7 mg/kg live weight, respectively (Surai, 2002; Sobolev et al., 2018). Feeding laboratory rats with SeO 3 2- and Se-yeast for 8 weeks manifested severe cardiotoxicity, hepatotoxicity, and splenomegaly in rats fed with SeO 3 2- whereas no such symptoms were exhibited in rats fed with the latter. Further histological examinations revealed that Se-yeast had
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greater deposition of Se. (Spallholz & Raftery, 1987). Clinical symptoms of Se poisoning in farm animals (cattle and sheep) are excessive salivation,
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garlic smell when exhaled, vomiting, loss of appetite, difficulty in breathing (shortness of
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breath), seizures, hoof deformation, lameness, diarrhea, alopecia (hair loss), bloating, loss of vision, exhaustion, paralysis and death, often from respiratory failure (Sobolev et al., 2018).
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Functions and tertiary enzymes structure are altered due to the formation of Se trisulfide
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complex (Se + Cys + coenzyme A). This could compromise the immune system due to the inability of the enzymes to catalyze activities, i.e., digestion, breathing, nutrient, absorption, etc.
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Moreover, Se toxicosis can lead to the production of methyl selenide (Se2-), a precursor for inducing oxidative stress on the system (Mézes & Balogh, 2009; Sobolev & Pacelja, 2016).
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A controlled clinical trial of testing a chemopreventive agent against prostate cancer progression by supplementing the diet of 24 men with either 1600 or 3200 µg/day of Se-yeast was carried
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out. The group fed with 3200 µg/day were reported to show symptoms such as garlic breath, brittle nails, and hair, stomach upset and dizziness unlike the former group (Reid et al., 2004). No side effects were documented when a similar study was replicated on hepatitis-B surfaceantigen positive patients (n=226) (Yu, Zhu, & Li, 1997). The latter researchers utilized 200 mg/d Se whereas the former supplemented with 3200/1600 mg/d. Other factors such as racial difference, age, health status, etc., could all be attributed to these findings. Rigorous studies are needed to elucidate the toxicity levels of Se taking into consideration the factors above (racial difference, age, health status). Table 3 shows the recommended daily Se intakes in some selected countries.
ACCEPTED MANUSCRIPT 7. Impact Se on gut microbiota. Microorganisms constitute the gut microbiota of every mammal and are essential in digestion, recycling of nutrients, and inhibiting colonization by pathogenic organisms (Guarner and Malagelada, 2003). Microbes residing in the gut are known to be actively involved in drug, Se, and vitamin metabolism (Boyle et al., 2006; Jia et al., 2008; Kasaikina et al., 2011b). Previous
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works have shown that Se administered as SeMet and other forms undergo a significant enterohepatic recycling, which involved the gut microflora (Xia et al., 1992; Krittaphol et al.,
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2009; Krittaphol, McDowell, Thomson, Mikov, & Fawcett, 2011a; Kasaikina et al., 2011; Saini
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& Tomar, 2017).
It was reported that Se could alter the composition of the gut microbiota, consequently
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influencing the selenoproteome and Se levels of the host. However, the composition of the diet
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coupled with other trace elements could influence the bacterial population in the gut (Kasaikina et al., 2011). Therefore, Se could impact commensal bacteria in the host that regulate nuclear
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factor-kB (NF- k B) and peroxisome-proliferator-activated receptor (PPARγ) which aids in the activation of immune cells during inflammatory bowel disease (IBD) (Nettleford & Prabhu,
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2018).
Gut microbiota contributes to the excretion of excess Se through the production of methyl Se
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compounds and elemental Se, thus protecting the host from selenosis (Krittaphol, McDowell, Thomson, Mikov, & Fawcett, 2011a). Probiotics like LAB strains improve the degradation of proteins in milk products, thus increasing the bioaccessibility of Se (Shen et al., 1996; Saini & Tomar, 2017). Enterococcus durans LAB18s was proposed as a potential probiotic that could be used for designing Se-enriched products. The highest percentage of bioaccessible organic Se was found in the fraction of total protein, followed by the fraction of polysaccharides and nucleic acids (Pieniz et al., 2017). The microbiota diversity could increase, thereby keeping the integrity of the gut microbiota intact due of Se bioavailability. Disruption of intestinal microbiota could lead to diseased states and
ACCEPTED MANUSCRIPT disorders (Vijay-Kumar et al., 2010; Kasaikina et al., 2011; Virgin & Todd, 2011). Motor control and anxiety behavior were reportedly correlated to host microbiota (Diaz Heijtz et al., 2011) In vitro probiotic tests confirms the ability of LAB to survive in the simulated gastric juice at pH 2 and could be exploited for the development of Se enriched functional foods (Se-enriched yogurt, kefir, beer, etc.) as a source of dietary Se (Saini, Tomar, Sangwan, & Bhushan, 2014; et al., 2014; Saini & Tomar, 2017). According to Saini et al. (2014), L. reuteri
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Pophaly
NCDC77 is a novel strain with greater potential to biotransform inorganic Se from a medium
(CH3 SeCys)
and
the
latter
transforms
to
highly
bioaccessible
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methylselenocysteine
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than the control bacteria. Additionally, it synthesizes selenocysteine (dimer of SeCys), and
methylselenol (efficient than SeMet) by the action of β-lyase and is considered as the most
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effective form of organic Se utilize in cancer therapy.
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In vitro and in vivo studies demonstrate that a variety of probiotic bacteria can metabolize SeMet and Na2 SeO 3 to produce methylated Se compounds. Probiotic mixtures significantly increased
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Se content in the liver and decreased them in the kidney after administration of SeMet compared to the control rats (Krittaphol et al., 2011b). In a similar study, an improved HPLC method was
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proposed for the quantification of L-SeMet in rat gut content suspensions prepared from the different parts of the intestine (jejunum, ileum, caecum, and colon). L-SeMet metabolism in the
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gut occurs mainly in the caecum because it houses microorganisms with high metabolic activity towards L-SeMet. Moreover, caecum is known to play a vital role in digestion and therefore contains a myriad of enzymes produced/secreted by the gut microbiota (Hall, Youngs, Keighley, 1992; Kurosawa, Ikeda, Sukemori, & Kurihara, 2005; Krittaphol et al., 2009). Further work is necessary to clarify into the biotransformation of L-SeMet by gut microflora (Krittaphol et al., 2009) about the bioavailability of Se. Lastly, in vivo and in vitro elucidation of Se metabolism pathway of LAB strains to get molecular and technological insights on prospective cultures and to model their response in fermented foods (Pophaly et al., 2014) need to be considered in future.
ACCEPTED MANUSCRIPT 8. Factors to consider when designing functional food 1. The enrichment/fortification must not alter the organoleptic properties (flavor, color, texture or odor) of the original food. 2. The active ingredient (Se) should demonstrate acceptable stabilities in the fortified foods. 3. The potential for any interactions that could occur between micronutrients in multiple
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fortified systems, as well as interactions between the active ingredient and food vehicle that may interfere with the metabolic uptake of the active ingredient, also need to be
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thoroughly examined.
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4. The addition of extra additives such as binders encapsulates, and stabilizers to improve the retention of the active ingredient should not require significant changes to existing
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technologies.
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5. The functional food must be well absorbed from the food vehicle at the level of consumption compatible with a healthy diet.
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6. The final cost of designing functional foods must not decrease the affordability of the food or increase the competitiveness with the unfortified alternative (Sangakkara, 2011).
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9. Conclusions and future trends
In developed nations, functional foods are considered to be and are ranked as one of the critical
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state policies. Adequate nutrition promotes growth and development, prevents diseases and increases productivity and life expectancy. Health risks associated with Se deficiency have affected many lives negatively and remains a threat to humanity. Therefore, enriching foods, which are consumed on a regular basis by a large portion of the population with Se, is the ultimate solution in combating the menace of Se deficiency.
Brewing yeast has the necessary
enzymes to biotransform inorganic Se (Na2 SeO 3 ) to organic form which is incorporated in the aged beverage. Se-enriched beverages are safe for consumption; hence, consumers should not worry about the issue with toxicity. S. cerevisiae was found to possess better biotransformation efficiency than the other yeast species. Selenized yeast and algae could be taken as supplements
ACCEPTED MANUSCRIPT or as food additives. However, the safety of Se-algae is not guaranteed because, recently, there have been some reports about MCs in Spirulina. Consumers should be careful about the source of Se-algae they are purchasing since the concentration of MCs depends on several factors, i.e. geographical location, strain, type of fertilization practice, etc. The public is again advised against Brazil nuts. It contains about 544 mcg of Se which can cause Se poisoning.
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Recent advances in the field of nutrigenomics are vital tools for investigating functional food. Deciphering the interactions between bioactive compounds (Se in this study) and proliferation of
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diseases based on an individual’s genetic profile is the ideal strategy to maximize health benefits
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and reduce disease risk from this food. The application of Nutrigenomics in the study of antiaging, obesity, cancer, etc. coupled with Se-enriched (functional food) is the best tool to
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understand the comprehensive benefits and side effects an individual gets from these foods.
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Thanks to Ventor and colleagues who completed the human genome sequence in 2001 (The Celera Genomics Sequencing Team, 2001), hence making the study of Se on individual genetics
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feasible.
The introduction of systems biology coupled with biotechnology is another vital tool in the
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future advancement of functional foods. Metabolic pathways could be engineered either to suppress the expression of genes responsible for the accumulation of excess Se in crops (i.e.,
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Brazil nuts) or to develop new strains of probiotics capable of secreting organic Se in the products. Nevertheless, this tool could also be used in developing genetically modified produce enriched with Se and other vital minerals, which could potentially improve the health of the consumer.
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Authors’ Contributions
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AP, NVD, KYM, and EFK have contributed to the study conception and design; AP searched
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for literature and wrote the manuscript; NVD, KYM, and EFK were responsible for the graphs, tables, and references. All authors have read and approved the final manuscript.
Acknowledgments
This work was financially supported by the Government of Russian Federation, Grant 08-08.
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ACCEPTED MANUSCRIPT Yan, L., & Spallholz, J. E. (1993). Generation of reactive oxygen species from the reaction of selenium compounds with thiols and mammary tumor cells. Biochemical Pharmacology, 45(2), 429–437 Yan, L., Reeves, P. G., & Johnson, L. K. (2010). Assessment of selenium bioavailability from naturally produced high-selenium soy foods in selenium-deficient rats. Journal of Trace Elements in Medicine and Biology, 24(4), 223–229.
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Yan, L., & Johnson, L. K. (2011). Selenium bioavailability from naturally produced high-selenium peas
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Yang, G., Yin, S., Zhou, R., Gu, L., Yan, B., Liu, Y., & Liu, Y. (1989b). Studies of safe maximal daily
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selenium intake in a seleniferous area in China. Part II. Relation between Se-intake and the
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manifestation of clinical signs and certain biochemical alterations in blood and urine. Journal of trace elements and electrolytes in health and disease, 3(3), 123–130.
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Yang, G., & Zhou, R. (1994). Further observations on the human maximum safe dietary selenium intake in a seleniferous area of China. Journal of trace elements and electrolytes in health and disease,
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8(3/4), 159–165.
Yang, L., Sturgeon, R. E., McSheehy, S., & Mester, Z. (2004). Comparison of extraction methods for
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quantitation of methionine and selenomethionine in yeast by species specific isotope dilution gas chromatography–mass spectrometry. Journal of Chromatography A, 1055(1/2), 177–184, Yin, H., Chen, Z., Gu, Z. & Han, Y. (2009). Optimization of natural fermentative medium for seleniumenriched yeast by D-optimal mixture design. LWT - Food Science and Technology, 42(1), 327– 331. Yoshida, M., Fukunaga, K., Tsuchita, H., & Yasumoto, K. (1999). An evaluation of the bioavailability of selenium in high-selenium yeast. Journal of Nutritional Science and Vitaminology, 45(1), 119– 128.
ACCEPTED MANUSCRIPT Yu, S. Y., Zhu, Y. J., & Li, W. G. (1997). Protective role of selenium against hepatitis B virus and primary liver cancer in Qidong. Biological Trace Element Research, 56(1), 117–124. Yuan, L., Yin, X., Zhu, Y., Li, F., Huang, Y., Liu, Y., & Lin, Z. (2012). Selenium in plants and soil, and selenosis in Enshi, China: Implication for selenium biofortification. In X. B. Yin, & L. X. Yuan (Eds.). Phytoremediation and biofortification: Two Sides of One Coin (pp. 7-31). Berlin:
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Springer. Zarea, H., Owlia, P., Vahidi, H., Khujin, M. H., & Ali Khamisabad, A. (2017). Yeast enriched with
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selenium: a promising source of selenomethionine and seleno-proteins. Trends in Peptide and
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Protein Sciences, 1(3), 130–134.
Zhang, Y., & Frankenberger, W.T. Jr. (2001). Speciation of selenium in plant water extracts by ion
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Total Environment, 269(1/3), 39–47.
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exchange chromatography-hydride generation atomic absorption spectrometry. Science of the
Zhang, B., Zhou, K., Zhang, J., Chen, Q., Liu, G., Shang, N., Qin, W., Li, P., & Lin, F. (2009). Accumulation
and
species
distribution
of
selenium in
Se-enriched
bacterial cells
of
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Bifidobacterium animalis 01. Food Chemistry, 115(2), 727−734.
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Zhou, N., Xiao, H., Li, T. K., Nur-E-Kamal, A., & Liu, L. F. (2003). DNA damage-mediated apoptosis induced by selenium compounds. Journal of Biological Chemistry, 278(32), 29532–29537.
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Zhu, Y. G., Pilon-Smits, E. A. H., Zhao, F. J., Williams, P. N., & Meharg, A. A. (2009). Selenium in higher plants: understanding mechanisms for biofortification and phytoremediation. Trends in Plant Science, 19(8), 436–442.
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Tables: Table 1. Total selenium in selenized beverages Se (IV) added
Se in selenized
(µg/mL)
beverages (µg/mL)
0.1
0.086±0.003a
1.0
0.61±0.08a
2.0
1.1±0.4a
10.0
6.0±0.7a
20.0
6.0±0.5a
30±5
50
15±1b 30±6b
30.6
150
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200 300
55±4 60±7
47±5b
30.8
115±8b
57.5
167±6b
55.6
300±10b
60.3
300±10b
29.7
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500
61±3
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43±5
30.3
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100
1000
% Se
Modified with permission from ref 4390651395301(Elsevier, 2011) and 4390660155828 (Elsevier, 2012). a
selenized-enriched beer after 12 days fermentation (Sánchez-Martínez et al., 2012)
b
selenized-enriched wine after 21 days fermentation (Pérez-Corona et al., 2011)
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Table 2. Different sources of functional foods/beverages containing selenium Reference
Se-enriched mushroom
Ogra, Ishiwata, Encinar, Lobinski, & Suzuki, 2004
Se-fortified yogurt
Achanta, Aryana, & Boeneke, 2007
Se-enriched eggs
Fisinin, 2007; Gajčević, Kralik, Has-Schön, & Pavić,
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Products
2009; Fisinin, Papazyan, & Surai, 2009; Bennett &
Hu, Xu, & Pan, 2001; Xu et al., 2003; Vodnar &
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Se-enriched tea
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Cheng, 2010
Socaciu, 2014
Lintschinger, Fuchs, Moser, Kuehnelt, & Goessler,
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2000 Se-biofortified apples
Wortmann & Daum, 2018
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Se-biofortified prickly pear
Tyrala, Borschel, & Jacobs, 1996
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Se-fortified infant formula Se-enriched fermented milk
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Se-enriched sprout
Alzate, Perez-Conde, Gutierrez, & Camara, 2010 Banuelos et al., 2011 IP & Lisk, 1993; Ip & Lisk, 1995, Lu et al., 1996
Se-biofortified onion
Adhikari, 2012; Golubkina et al., 2016
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Se-enriched garlic (Se-garlic)
Se-biofortified broccoli
Pedrero, Elvira, Camára, & Madrid, 2007; Adhikari, 2012
Columbus Egg
De Meester et al., 2000
Se-enriched beef
Lawler, Taylor, Finley, & Caton, 2004; Lee et al., 2006; Taylor et al., 2008
Se-enriched wine
Pérez-Corona et al., 2011
Se-enriched beer
Sánchez-Martínez et al., 2012
Se-enriched kvass
Muravyov, 2018
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Liangcai, 2015; Wave et al., 2017
Se-enriched bread
Sarudi, Lassu-Merényi, Rumi, & Kelemen, 1997; Bryszewska et al., 2007; Hart et al., 2011; Lazo‐ Vélez, Chávez‐ Santoscoy, Serna‐ Saldivar, 2014 Yan & Johnson, 2011
Se-biofortified shallot
Ogra, Ishiwata, Iwashita, & Suzuki, 2005
Se-biofortified pumpkin
Smrekolj, Stibilj, Kreft, & Kápolna, 2005
Se-biofortified dill
Cankur, Yathavakilla, & Caruso, 2006
Se-biofortified chicory
Mazej, Falnoga, Veber, & Stibilj, 2006
Se-biofortified radish
Ogra et al., 2007
Se-biofortified ramp
Whanger, Ip, Polan, Uden, & Welbaum, 2000
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Se-fortified oat
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Table 3. Current recommended selenium intakes for adults (µg/d) Male
Females
USA and Canada, 2000
55
55
Australia, 1990
85
70
Belgium, 2000
70
70
30-70
30-70
55
55
60
50
55
55
55-60
45
65
55
50
40
75
60
40
30
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Countries
Germany, Austria, Switzerland, 2000
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EC Scientific Committee on Food, 2003
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France, 1996 Italy, 1999
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Japan, 1999
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New Zealand and Australia (proposed levels) Nordic countries, 1996
World
Health
Organization/Food
and
Agriculture
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Organization/International
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UK (Committee on Medical Aspects of Food Policy, 1991
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Atomic Energy Agency, 1996
Adapted with permission from ref 4392370266741 (Cambridge University Press, 2006) (EC Scientific Committee on Food, 2003; Thomson, 2004; Rayman, 2004).
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Figures:
Figure 1. The process of brewing Se-enriched beer.
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Figure 2. Production processes of selenized kvass.
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Figure 3. Transformation and bioaccumulation of Se in algae. 1) Selenium uptake via sulfate, phosphate and silicon transporters. 2) Selenium/sulfur assimilation pathway: APSe: adenosine
(seleno)methionine;
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phosphoselenate; GSH: glutathione; OAS: O-acetylserine; (Se)Cys: (seleno)cysteine; (Se)Met: MMT:
dimethyl(di)selenide
methyltransferase; (volatile);
SMT:
DMSeP: selenocysteine
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dimethylselenoproprionate;DM(D)Se:
methylmethionine
methyltransferase. 3) SeCys is incorporated in essential Se-proteins that play a key role in ROS scavenging and defense systems. 4) However, if intracellular Se concentration is high, Se amino acids can also be incorporated into other proteins non-specifically, thus producing malformed structures (Schiavon et al., 2017). Modified with permission from ref 4390660547884 (Elsevier, 2017)
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Figure 4. The production of Se-enriched Tom-Brown flour.
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Figure 5. Comparing toxicity levels of the various form of selenium (sodium selenite and selenium-enriched yeast). Modified with permission from ref 4392380281814 (Cambridge University Press, 2006).
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Graphical abstract
ACCEPTED MANUSCRIPT Highlights
Organic selenium (Se) is required to sustain proper health in both animals and humans due to its linkage with various biological functions in the immune system.
Some regions around the world are naturally deficient in soil Se whilst others are becoming toxic. Organic Se is more bioavailable and less toxic than the inorganic form.
Studies have shown that the consumption of Se enriched food and supplements leads to
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Supplementing wort with inorganic Se resulted in brewing selenized beverages (beer and
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Kvass).
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the decrease of selenosis.
Figure 1
Figure 2
Figure 3
Figure 4
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Parise Adadi, Nadezhda V. Barakova, Kirill Y. Muravyov, Elena F. Krivoshapkina PII: DOI: Reference:
S0963-9969(18)30914-1 https://doi.org/10.1016/j.foodres.2018.11.029 FRIN 8092
To appear in:
Food Research International
Received date: Revised date: Accepted date:
23 July 2018 15 October 2018 15 November 2018
Please cite this article as: Parise Adadi, Nadezhda V. Barakova, Kirill Y. Muravyov, Elena F. Krivoshapkina , Designing selenium functional foods and beverages: A review. Frin (2018), https://doi.org/10.1016/j.foodres.2018.11.029
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ACCEPTED MANUSCRIPT Designing selenium functional foods and beverages: A review
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Parise Adadi*, Nadezhda V. Barakova, Kirill Y. Muravyov, Elena F. Krivoshapkina
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ITMO University, Lomonosova Street 9, 191002, St. Petersburg, Russian Federation
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*Corresponding author
ITMO University,
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Lomonosova Street 9, 191002,
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Parise Adadi
St. Petersburg, Russian Federation
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E-mail address: [email protected]; adadi_parise@scamt- itmo.ru.
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Tel: +79817511640
ACCEPTED MANUSCRIPT ABSTRACT A functional food is any food that has (a) specific nutrient(s) added to it for a specific functional purpose. Selenium (Se) is a metalloid that belongs to group 16 of the periodic table. It may be obtained from myriad sources like soil, water, and living organisms. Se is required to sustain proper health in both animals and humans due to its linkage with various biological functions in
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the immune system. Nature has made it impossible for us to obtain sufficient Se from the diet since some regions across the globe have been designated as Se-deficient while others are
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becoming Se-toxic. Se deficiency is associated with a compromised immune system and
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increased susceptibility to various diseases. Therefore, designing Se functional foods and supplements is the way forward in curbing the menace mentioned above, since geographical
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location will not have any effect on the Se content of these foods. Brewing yeast has the
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necessary enzymes to biotransform inorganic Se (Na 2 SeO 3 ) to its bioactive organic form, which is incorporated in the aged selenized beer. S. cerevisiae was found to have better
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biotransformation efficiency than other yeast species. A traditional Slavic beverage, selenized kvass, was brewed using rye grains soaked and germinated in solutions of Na 2 SeO 3 . Fruit yeasts
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(Lesaffre) and Evitalia were utilized as the starter cultures. A Se enriched solution was extracted from Se-biofortified pak-choi cabbage and incorporated into the wort before fermentation.
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Selenized yeast and algae can be taken as supplements or as food additives. There have been some reports about microcystins (MCs) in Spirulina. Therefore, the safety of Se-algae is not guaranteed. Pasteurized dried selenized supplements (yeast and algae) were proposed in formulating Selenized enriched Tom-brown. This review seeks to propose some possible foods that could be enriched with Se. A large portion of the population adequately consumes these proposed foods on a regular basis hence the target goal would be a success.
Keywords: Selenized beer, Slavic foods (Kvass), Yeast, Algae, Selenosis, Anti-cancer effects
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Abbreviations. FOSHU – foods for specified health use; FUFOSE – functional food science in Europe; SHF – specific health promoting food; TDI – tolerable daily intake; MCs – microcystins; SeCys – selenocysteine;
SeMeSeCys
–
selenomethyl-selenocysteine;
SeMet
–
selenomethionine;
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NaHSeO 3 – sodium hydroselenite; kDa – Kilo Dalton; Se-yeast – selenized-yeast; Se-algae – selenized algae; Se – Selenium; ICP-MS – liquid chromatography hyphenated to an inductively
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coupled plasma mass spectrometer Selenium; MHLW – Ministry of health, labor and welfare;
inductively
coupled
plasma
mass
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SOD – superoxide dismutase; CAT – catalase; GSH-Px – glutathione peroxidase; ICP-MS – spectrometry;
HPLC
–
High
Performance
Liquid
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Chromatography; MMeSe – monomethyl form of Se; DMeSe – dimethyl form of Se; TMeSe –
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trimethyl form of Se; SDS – Sodium dodecyl sulfate.
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1. Introduction In the last few decades, the world has experienced an increase in the cost of healthcare and people have found this alarming. Hence, they are searching for ways on how to stay healthy in order to avoid excessive spending on healthcare. Presently, the perception of food has changed
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from just consuming/drinking something to fill the stomach but rather eating to benefit from the nutrition, which enhances the physiological functioning of the system. At about this time the
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Japanese approved the process for functional foods called Foods for Specified Health Use
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(FOSHU) in the 1980s (Arai, 1996). There is no universal definition for functional foods as they vary across countries (The Japanese ([FOSHU]), European (Functional Food Science in Europe
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[FUFOSE]), and Dutch (Specific Health Promoting Food [SHF])). For instance, FUFOSE
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defined a functional food as “food that beneficially affects one or more target functions in the body, beyond adequate nutritional effects, in a way that is relevant to either an improved state of health and well-being and reduction of risk of disease. It is part of a normal food pattern. It is not
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a pill, a capsule or any form of dietary supplement.” The Academy of Nutrition and Dietetics
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sees it as “whole food along with fortified, enriched, or enhanced foods that have a potentially beneficial effect on health when consumed as part of a varied diet on a regular basis at effective
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levels.” Conversely, FOSHU described it as foods containing an ingredient with functions for health and officially approved to claim its physiological effects on the human body (Academy of Nutrition and Dietetics, 2013). Aryee & Boye (2015) have documented various health benefits of functional foods.
According to Cocks, Wrigley, Chicarelli-Robinson, & Smith (1995), herbs, algae, and microorganisms are rich sources of bioactive ingredients used in designing functional foods. Functional foods must remain foods (should not be pills or capsules) mostly composed of bulk ingredients (fruits, beverages, vegetables, cereals, nuts, milk, and milk-based products), and they
ACCEPTED MANUSCRIPT must demonstrate their effects in amounts that can be expected for normal consumption patterns (Sandhu & Sra, 2014). The demand for functional foods has increased over the past decades, resulting in the exploitation of this need by entrepreneurs who have established multibillion-dollar companies mainly invested in designing foods to satisfy consumers’ needs. These companies have seen
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significant growth, mostly in developed countries (United States, Canada, Denmark, Ireland, etc.).
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Selenium (Se) is a metalloid that belongs to group 16 of the Mendeleev's periodic table with
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physicochemical properties similar to sulfur due to their proximity (ionic radius) on the periodic table. The biological and toxicological effects of Se strongly depend on its chemical form,
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therefore the organic form is preferable to the inorganic form due to its bioavailability (Bodnar,
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Konieczka, & Namiesnik, 2012; Gupta & Gupta, 2016). Se is an essential nutrient in human life and participates in some biochemical reactions in the
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body. As reported by Oldfield (1999), Se is directly linked to a spectrum of biological activity. It exhibits similar antioxidant properties (act like superhero nutrients) to lycopene (Adadi,
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Barakova, & Krivoshapkina, 2018), thus protecting the body against damaging free radicals. Se act as a cofactor for triiodothyronine deiodinases, an important enzyme involved in thyroid
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hormone metabolism, thereby affecting iodine status and consequently preventing goiters (Arnaud et al., 2001; Erdenetsogt, Golubkina, Nadegkin, Monhoo, & Batjargal, 2014). It also acts as a cofactor for various enzymes, for example, glutathione peroxidase (GSH-Px), superoxide dismutase (SOD), and catalase (CAT), which shield the cells against damage triggered by free radicals and lipoperoxides. (Thompson & Scott, 1969; Newberne & Suphakarn, 1983). Health disorders such as infertility, cardiovascular diseases and cancer (oxidative stress) are associated with the insufficient intake of Se (Broadley et al., 2006). Human in vitro studies suggest that supplementing the diet with Se decreased the incidence of cancer (Combs, 2005; De Martino, Filomeni, Aquilano, Ciriolo, & Rotilio, 2006).
ACCEPTED MANUSCRIPT It is estimated that the diets of as many as 1 billion people might lack sufficient Se for their wellbeing (Combs, 2001; Fairweather-Tait, Collings, & Hurst, 2011; Joy et al., 2014; Stoffaneller & Morse, 2015). Nevertheless, Se deficiency and toxicity remain a problem in many human populations (National Research Council, 1983). Plant products are the major source of Se. Human low dietary Se intake is associated with crops
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cultivated in soils low in Se (Broadley et al., 2006; White & Broadley, 2009; Chilimba et al., 2011; Fairweather-Tait et al., 2011; Rayman, 2012; Fordyce, 2013; Joy et al., 2015).
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Consequently, animal-based food could serve as an alternate source since animals can
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accumulate Se from Se-supplemented feed or pastures fertilized with Se fertilizers. Meat, sausages, eggs, milk, apples (Se-biofortified apples), dairy products, fish are the primary sources
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of Se in the human diet, although this varies across regions (Meyer, Heerdegen, Schenkel,
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Dänicke, & Flachowsky, 2004; Rayman, 2008; Rayman, 2012; Wortmann, Enneking & Daum, 2018). Zhu, Pilon-Smits, Zhao, Williams, & Meharg (2009) found that some regions around the
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world are Se-deficient while others are becoming Se-toxic due to natural and anthropogenic activities.
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There is a direct correlation between soil Se content and the Se dose in products obtained from such soils (Finley, Matthys, Shuler, & Korynta, 1996). The soils in Sverdlovsk, Chelyabinsk,
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Tyumen, Novosibirsk, Irkutsk, Chita, Magadan, Amur, Krasnoyarsk, Khabarovsk, Primorsky Krai, Sakha-Yakutia, Komi, Mari-El Republic, Karelia, St. Petersburg, the Upper Volga region, Yaroslavl region and Udmurt Republic are naturally deficient in Se (Golubkina & Alfthan, 1999). This predisposes the inhabitants of these regions to mild selenosis with symptoms ranging from dermatitis, cracking of nails, hair loss and garlicky breath (due to exhalation of dimethylselenide). Severe selenosis can lead to acute respiratory distress, myocardial infarction and renal failure (White, 2015). As reported by Dhillon & Dhillon (2003) and Fordyce, (2013), animals also suffer from this disease, with symptoms ranging from
hair loss, garlicky breath,
hoof deformation (in horse, cattle, donkey), abnormal posture, lack of vitality, growth disorder,
ACCEPTED MANUSCRIPT anorexia, diarrhea, reduced reproductive performance, fetal deformities and respiratory failure due to insufficient intake of Se. Se deficiency is not only bound to Russia but other areas around the world (Zhu et al., 2009; Yuan et al., 2012). Se fertilizers were formulated to enable farmers to fertilize their crops (biofortification). Products from these farms were found to contain more Se than the control
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farms (Finley et al., 1996). The problem with toxicity and volatility of Se salts at high temperatures when applied also
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surface and remains a challenge to farmers. However, the problem with toxicity could be curbed
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by planting specialized crops, namely Se accumulators (e.g., medicinal melon) in the contaminated soil (White, 2015; Gupta & Gupta, 2016). For these reasons, Se-enriched foods
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and supplements have been proposed to circumvent the problems related to Se deficiency.
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Therefore, some technologies for designing Se-enriched functional foods, a mechanism for toxicities and health concern were discussed. The impact of Se on gut microbiota, its
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bioavailability, as well as the factors to consider when adopting the techniques mentioned above were also pondered in this manuscript.
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2. Production of beverages enriched with selenium Beverages are drinks prepared by the combination of two or more materials (i.e., water, sugar,
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colors, additives, yeast, etc.) for human consumption except for the regular water we drink every day. They are divided into two types (i) alcoholic (ii) non- alcoholic beverages. Beer, wine, and palm wine, etc., are categorized as alcoholic beverages while tea, kvass and fruit drinks are grouped under non-alcoholic beverages. These classifications vary according to the standard of each country. Beverages are not just for quenching thirst but also provide the consumers with energy, vitamins, minerals (Fellows & Hampton, 1992) and other vital bioactive substances required for the normal physiological function systems. In 2012, about 923 billion liters of commercialized beverages were sold across the world, with consumption in the United of States of America, Germany, and Brazil estimated at 182.2, 51.1,
ACCEPTED MANUSCRIPT 62.2 billion liters of beverages respectively in 2013 fiscal year alone (Euromonitor International, 2012; Madi, Castro, & Wallis, 2016). In producing these beverages, single or mixed starter cultures are usually employed. These cultures possess the vital enzymes to biotransform inorganic Se to more bioactive organic forms. As reported by Nelson, (2005), beer is one of the world’s most known and widely consumed drink after tea and water. Hence, enriching this
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product with Se could be an excellent intervention in combating deficiency of this micronutrient. While alcohol consumption is prohibited in Muslim countries, non-alcoholic beverages could
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also be designed as alternate conveyors to compensate the Se deficiencies in those countries.
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2.1 Supplementation of Se during brewing
Brewing yeast - Saccharomyces cerevisiae, and lactic acid bacteria (LAB) have shown the
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ability to transform inorganic Se to the organic form (safer, preferred and highly bioactive)
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(Schrauzer, 2000; Alzate et al., 2007; Alzate et al., 2008). This has drawn the attention of brewers to look into this novel technology in order to brew Se enriched beers.
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Metabolizing low bioavailable Se compounds (inorganic Se) into their high bioavailable forms (organic Se) by a plant, yeast, bacteria was reported (Kurek et al., 2016). Sodium selenite
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(Na2 SeO 3 ) was supplemented in media cultured with Bifidobacterium animalis (Zhang et al., 2009) as probiotic bacteria used in producing Se-enrich products. The single infusion method of
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mashing could be used, where the malt is mixed with distilled water in the mash tun and heated at 45 °C for 30 minutes. The temperature is then increased to 54 °C and then 62 °C with step times of 40 and 45 min respectively. After these step times, the temperature will once, again be increased to 72 °C for 55 min. Finally, the temperature is increased to 78 °C for 10 minutes. Filtration is carried out while 1 L distilled water is used for the sparging process. The wort is then boiled at 100 °C for 90 min, hops is then added 80 min before the end of the boiling time. The wort is then allowed to cool to the desired temperature (20 o C) by either placing the boiled wort under running cold water or using a chilled sterilized copper wort chiller. The wort is then transferred to sterilized fermentation vessels equipped with airlock bubblers (Adadi, Kovaleva,
ACCEPTED MANUSCRIPT Glukhareva, Shatunova, & Petrov, 2017; Adadi, Kovaleva, Glukhareva, & Barakova, 2018) where a different concentration of Na2 SeO 3 is added (Sánchez-Martínez et al., 2012). Even though Na2 SeO 3 could be used as a precursor for the formation of selenoproteins in beer, it cannot be stored for future use (Alfthan, Aro, Arvilommi, & Huttunen, 1991). The flow chart of brewing Se enriched beer is depicted in figure 1. The yeast is then pitched while the fermentation
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kinetics is studied. Fermentation is then carried out at a temperature of 25 °C for 5-12 days depending on the type of Se-enriched beer brewed.
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The total Se and Se species could be determined using inductively coupled plasma mass
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spectrometry (ICP-MS) and liquid chromatography hyphenated to an inductively coupled plasma mass spectrometer (HPLC–ICP-MS) respectively. The details are documented in Perez-Corona
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et al. (2011) and Sánchez-Martínez et al. (2012).
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The fermentation process is not altered due to the addition of Se in the wort or must (PerezCorona et al., 2011). Se does not alter the sugars and other nutrients vital for the yeast, hence the
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fermentation process will proceed without any problems. Yeasts are excellent accumulators of Se and other trace elements (Schrauzer, 2006; Kieliszek & 2013).
Saccharomyces
cerevisiae
and
Lactobacillus have the enzyme for
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Błażejak,
biotransformation of inorganic Se to organic form and they are widely utilized in brewing
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beverages (Sánchez-Martínez et al., 2012). Much research is devoted to the study of intracellular accumulation and metabolism of Se, mainly by yeast and bacteria (Ponce de León, Bayon, Paquin, & Caruso, 2002; Gharieb & Gadd 2004; Pieniz, Andreazza, Mann, Camargo, & Brandelli, 2017). Kieliszeket, Błażejak, Gientka, & Bzducha-Wróbe (2015) tried to decipher the accumulation and transformation of Se in yeast cells. The added Na2 SeO 3 is metabolized in two possible pathways, i.e. by methylation coupled with the reduction of Se and direct Se incorporation or bind to proteins in which Se replaces sulfur mainly by the amino acids cysteine and methionine (Fairweather-Tait et al., 2010; Kieliszek &
ACCEPTED MANUSCRIPT B1azejak, 2013). Kieliszek & B1azejak (2013), in their review, stated that selenates (SeO4 -2 ) are probably reduced to selenites (SeO 3 -2 ), elemental Se (Se0 ), and selenides (Se2-). In the presence of
yeast,
glutathione
and
adenosine-5-triphosphate
are
transformed
to
adenosine-5-
selenophosphate via enzymatic activation (i.e., Sul1 and Sul2 sulfate permeases, ATP sulfurylase, PAPSe reductase) in which SeO 4 -2 is converted to SeO 3 -2 . Other forms of Se such as
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monomethyl (monomethyl form of selenium (MMeSe), dimethyl (dimethyl form of Se (DMeSe), and trimethyl (trimethyl form of Se (TMeSe) can be obtained from non-methylated form, which
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is less toxic than other forms of Se (Pyrzynska, 1996; Bánszky, Simonics, & Maráz, 2003;
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Mapelli, Hillestrøm, Patil, Larsen, & Olsson, 2012; Kieliszek et al., 2015; Kieliszek & B1azejak, 2013). SeO 3 -2 reacts further with glutathione to form selenodiglutathione (GS-Se-SG), an
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oxidized form of glutathione (GSSG) (hazardous compound). GSSG is detoxified to the reduced
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form (GSH) by glutathione reductase in the vacuole. Glutathionyselenol (GS-Se-H) and hydrogen selenide (H2 Se) are formed because of further spontaneous metabolism of GS-Se-SG
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couple with GSSG. Via passive transport, H2 Se is secreted (through the vacuolar membrane) out into the cytoplasm of the cell. The activities of superoxide dismutase further synthesized
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glutathionylselenol into products (elemental Se and glutathione) (Tarze et al., 2007; Lazard et al. 2011; Mapelli, Hillestrøm, Kapolna, Larsen, & Olsson, 2011; Kieliszek et al., 2015). H2 Se is
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highly considered as a potent reducing agent along with glutathionyselenol (GS-Se-H), glutathione, and cysteine molecules (Kieliszek et al., 2015). H2 Se act as a precursor in the formation of organic compounds, including selenoamino acids where it is bound to O-acetyl homoserine (O-Ac-HSer) involving homocysteine synthase. Selenohomocysteine (SeHCys) and acetic acids are then synthesized (homoselenocysteine biosynthesis). The reaction is further branched into selenocystathionine or selenomethionine (SeMet) synthesis. SeHCys is bound to serine, which is then catalyzed by cystathionine βsynthase, forming selenocystathionine and water as a by-product. SeMet is substituted with formethionine because it easily acylates Met-tRNA which leads to the accumulation of
ACCEPTED MANUSCRIPT methionine-requiring proteins (The reaction is catalyzed by homocysteine methyltransferase). Another possible route is via trans-sulfuration mechanism, which is catalyzed by γ-lyase into selenocysteine (SeCys) and H2 Se (Pedrero & Madrid, 2009; Mapelli et al. 2011; Kitajima & Chiba 2013). The biosynthesis of SeCys in yeast cells is well elucidated in the following literature (Xu et al., 2007; Squires & Berry, 2008; Allmang, Wurth, & Krol, 2009; Turanov et
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al., 2011). Na2 SeO 3 reacts with reducing sugars in wort, forming the red elemental Se, which is toxic.
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(Mapelli et al., 2011). This toxic compound could end up in the Se-enriched beer. The ability of
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yeast to bind sulfur onto its cell wall is of interest to brewers. Sulfur act as a precursor for the formation of unpleasant flavors in some beers, which leads to loss of revenue since such
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products are poorly patronized.
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Rhodotorula mucilaginosa-13B could be used in cleaning water with a high Se dose due to its ability to biotransform and accumulate Se on its structures (Ruocco, Chan, Hanson, & Church,
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2014). S. uvarum can biotransform inorganic Se to organic form when pitched in nutrient-rich wort. This theory was confirmed in the work of Marinescu, Stoicescu, & Teodorof (2011) where
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they reported that Se became organically bound to the yeast when brewery wastewater was used as the medium for cultivation.
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It was reported by Sánchez-Martínez et al. (2012) that increasing the Se concentration in wort resulted in an increase in the percentage of Se in beer. However, above 10 µg/mL, the quantity remained the same. This result also depends on the kind of yeast (top or bottom fermenting) used. Thus, at a certain threshold, the yeast could no longer tolerate the toxicity of the incorporated Se. Therefore, this threshold should be deciphered so that brewers can manage and produce a better beer. This could also avoid brewing beer contaminated with toxic Se. Zare and
coworkers
(2017) reported that among the numerous yeasts screened for
biotransformation of inorganic Se to it bioavailable form, only S5, and S18 were found to yield good SeMet content (above 2500 ppm organic Se). Nevertheless, S18 isolate showed the
ACCEPTED MANUSCRIPT maximum biomass production and SeMet accumulation (2655 ppm) and (3.73 g/L) when exposed to Se at 25 mg/L concentrations. Pérez-Corona et al. (2011) reported the biotransformation of inorganic Se by S. cerevisiae and S. bayanus when they produced white wine under laboratory-scale. It was observed that the viability of cells in the must (must and inorganic Se) were lower compared to the control (YEPD
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medium) experiment, though this did not affect the entire fermentation process. The addition of 200–500 µg of SeO3 2- in the must resulted in the production of Se-enriched wine (55–60%, Table
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1). This percentage starts to decline (by approximately 30%) when the amount of SeO 3 2-
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increases, hence the maximum quantity of SeO 3 2- to incorporate in any beverage should be in the range between 200–500 µg. A scale-up experiment is warranted in order to validate the findings S. cerevisiae has higher biotransformation efficiency than the S. bayanus. Nevertheless,
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above.
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screening for Se toxicity level (tolerance level) of various yeast is a good initiative in selecting efficient strains for producing Se-enriched beverages with good sensorial properties and high
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acceptance levels. Since beer is one of the most consumed beverages in the world, Se beer can be considered an adequate source of Se for people deficient in Se.
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2.2. Production of selenized kvass
Kvass was invented by the traditional Slavs and is a favorite low alcoholic beverage in the
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Eastern European countries (especially in Russia and the Ukraine) and the former Soviet republics. It is produced from either rye/wheat bread (dried). The hue varies (color of bread to dark black) due to the utilization of different raw materials in the production process. By the Russian standard, the beverage is categorized as a non-alcoholic drink (0.5–1.0%). The demand for different flavors in kvass has caused producers to start adding fruits and other flavor enhancing substances (Hornsey, 2003; Sandor, 2003; Volhina, 2011; GOST, 2005). The entire production process of selenized kvass is summarized in figure 2. Rye grains were soaked and germinated in solutions of Na2 SeO 3 at room temperature for 4 days. The germinated
ACCEPTED MANUSCRIPT grains were then kilned in an oven to halt the enzyme activities. The enzymatic activities in the grains are thought to biotransform the absorbed SeO 3 2-. The malted rye grains were dried and weighed before crushing in a roller mill. The milled grains were then sieved through a screen with 1 mm openings (determined by weighing after milling) for better extraction during mashing.
Milled grains (500 g) was mixed with potable water and
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heated at 58 °C for 20 minutes. The ratio of the malt and water was 1:2.5. The temperature was increased to 64 °C and 72 °C with step times of 60 and 120 mins respectively. The temperature
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was finally increased to 78 °C for 10 mins. Filtration was carried with the aid of a cheesecloth o
Brix for the control and the Se-
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and 3.51 L (80 °C) of water was used for the sparging. The enrich wort was 11.7 and 12.4 respectively.
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Dry Se-biofortified pak-choi cabbage (3.69 g) (the concentration of the Se was 250 μg/kg) was
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ground to the size of 05.2 mm, water was then added at a ratio of 1:12. Enzyme preparations were made to enhance the extraction.
Distizim-protacid extra (manufactured by Erbsloh,
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Germany) and Viscostar (Novozymes manufacturer) had proteolytic and cellulolytic activities on the sample. The dose of enzymatic preparations was 1% per 1 kg of raw material. The
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temperature of the mixture (enzymes + sample) was raised to 55°C and held for 24 hours, the solution was filtered, and the Se concentration of the extract (filtrate) determined. The Se-
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enriched extract was poured into the wort. Fruit yeasts (Lesaffre) were reactivated before pitching for reparation of vital functions in stored yeasts, together with LAB Evitalia (1.25 g yeasts, 25 mg bacterial culture).
Fermentation lasted for 24 hours at 30 °C. Finally, the selenized kvass underwent pasteurization for 20 minutes at 80 °C (Antoņenko, Duma, Kreicbergs, & Kunkulberga, 2016; Muravyov, Barakova, Hamsters, & Panova, 2018a; Muravyov, Barakova, Hamsters, & Panova, 2018b; Muravyov, 2018).
It was allowed to cool before bottling followed by sensorial analysis
(selenized kvass). The Se-enriched extract could have been added to the green beverage before conditioning and not prior to fermentation. Quantification of Se in rye and the final products
ACCEPTED MANUSCRIPT were done according to Russian standard (GOST, 2008). Other techniques could be applied as well. The results revealed that selenized kvass produced from rye grains soaked and germinated in solutions of Na2 SeO 3 had the higher content of Se (51.1 μg/kg) when compared to the control experiment. However, the alcohol content (0.7%) was more than the control (0.5%), indicating that Se influenced the enzymatic activities in the malt. According to Antoņenko et al. (2016), the
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addition of Se significantly increased the activity of amylases. This might have increased the sugar content in wort, explaining the higher alcohol percentage recorded in the selenized kvass.
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3. Nutritional supplements (selenized-yeast (Se-yeast) and algae (Se-algae))
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Selenium-enriched supplements are ideal intervention to increase the Se intake for a large population, thus minimizing the risk of diseases and deficiencies associated with Se.
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The active groups of proteins, phospholipids, or polysaccharides of yeast cell wall acts as the
composed
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biological binder for Se (Kieliszek et al., 2015). According to the literature, the cell wall is of polysaccharides (85%) and
proteins (15%). Further examination of the
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polysaccharide revealed that it included glucose (80–90%), mannose (10–20%), and Nacetylglucosamine (1–2%) (Klis, Boorsma, & De Groot, 2006).
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The architecture of the cell wall is composed of inner layers of the carbohydrate polymer β-(1,3) glucan (30–45%) and the outer layers of mannoproteins (30–50% of the cell wall). Additionally,
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β-(1,6) glucan (8–18%) and chitin amount to 1-2% of the total composition of the cell wall (Klis et al., 2006; Saluk-Juszczak, Królewska, & Wachowicz, 2010; Levin & Moran 2011; Orlean, 2012).
Se-yeast is the product of the aerobic fermentation of various yeast species in a Se-enriched medium (Rayman, 2004). Yeast can bioaccumulate up to 3000 µg/g of organic Se (Schrauzer, 2006) which depends on the type of media, the conditions, and concentration of the media constituents. Inorganic Se is transformed into SeMet, which is the main selenoamino acid found in the enzymatic hydrolysis of yeast (Ponce de León et al., 2002; Rayman, 2004; Reyes et al., 2004;
ACCEPTED MANUSCRIPT Schrauzer, 2006). S. cerevisiae is one of the best organisms that can biotransform inorganic Se to the more bioactive SeMet form (Zare et al., 2017). Under stress conditions (high temperature, fluctuation in pH, etc.), yeast can convert inorganic Se into organic Se (SeMet) which can be accumulated on the cell wall. Increasing the scale of production of Se-enriched yeast is the last step in order to meet human
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demands for SeMet (Yin, Chen, Gu, & Han, 2009; Sanchez et al., 2012; Bierla, Szpunar, Yiannikouris, & Lobinski, 2012; Rajashree & Muthukumar, 2013b). However, green (clean)
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methods should be adhered to when fulfilling this goal in order to avoid cultivating yeast
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contaminated with heavy metals and other naturally occurring yeast strains. Bioaccumulation of SeMet (300 to 2200 ppm) was achieved with yeast cells using a synthetic medium. The yield was
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increased by supplementing the culture medium with Na2 SeO 3 (Suhajda et al., 2000; Antoneta,
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Marinescu, & Teodorof, 2011; Muthukumar & Rajashree, 2013; Zare et al., 2017). Many industries across the world have now mastered the large-scale production of Se-yeast. Its
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relatively low production cost makes it a promising and attractive venture for the food and pharmaceutical industries. A mathematical model was applied to study the effects of culture
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conditions (temperature, fermentation time, initial pH value, shaking speed, as well as time and concentration of Se, added to medium) on the bioaccumulation of Se in yeast. The total Se
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accumulation and organic Se formation ranges from 107.9 to 287.6 mg/kg and 93.27 to 269.05 mg/kg, respectively and the best conditions to obtain these results were: Se concentration, 25 μg/mL; addition time of Se source, 9 h; inoculums size, 30 g/L; shaking rate, 130 rpm; fermentation time, 48 h; temperature, 28 °C; and pH, 5.8 that have maximum Se incorporation (Esmaeili, Khosravi-Darani, Pourahmad, & Komeili, 2012). Se-yeast is the most studied Se-enriched product. However, Se species identification and quantification in this matrix are still a challenge (Pedrero & Madrid, 2009). Recent development has helped to solve this shortfall (Méndez, González, & Sanz-Medel, 2000; McSheehy, Pannier, Szpunar, Potin-Gautir, & Lobinski, 2002; Hinojosa-Reyes, Ruiz-Encinar, Marchante-Gayun,
ACCEPTED MANUSCRIPT Garcia-Alonso, & Sanz-Medel, 2006). Yeast Candida utilis ATCC 9950 was found to secrete Se in culture media which is triggered by enzymes associated with intracellular transport of Se. This leads to bioaccumulation of organic Se in the cell wall, which is a good source of dietary Se. Electrophoresis of the medium revealed protein fractions of yeasts obtained from media supplemented with Se, the presence of new protein fractions characterized by molecular weights
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of 60, 22, 18, and 15 kDa (Kieliszek et al., 2015). However, high accumulation of Se in S. cerevisiae yeast cells occurred in protein fractions above 75 kDa (Bryszewska, Pęczkowska,
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Rudzi´nski, Diowksz, & Ambroziak, 2000). The biosynthesis of selenoproteins in yeast cells is
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induced due to the presence of Se in the medium (Kieliszek et al., 2015), which are the bioaccessible form of Se. Se-yeast could be utilized in formulating different kind of foods i.e. ice
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cream, porridge, drinks, etc.
Se-enrich medium.
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According to Rayman (2004), Se-enriched yeast cream is the product after culturing yeast in a Pasteurization is then carried out, thereby killing the yeast, followed by
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spray drying. Dried Se-yeast can be used in formulating foods like ice creams and cereal basedcomplementary foods. Stabnikova et al. (2008) added a solution of sodium hydroselenite
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(NaHSeO 3 ) with a concentration of 100 µg Se/mL to the sterile media utilized for incubating S. cerevisiae. The growth (kinetics) and influence of Se on yeast biomass at different
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concentrations were studied. It was established that the growth was inhibited due to the addition of NaHSeO 3 to the medium, and resulted in the accumulation of elemental Se, which is not suitable for human consumption. The specific growth rates declined (0.146 to 0.073 L/h) when the Se concentration increased (from 2 to 12 µg/mL). The obtained Se-enriched yeast was used to bake Se-enriched bread. Low fermentative activities of yeast and longtime of the dough pellet to float up (rise power) were some of the problems faced by the authors. Nevertheless, they curb these problems by re-suspending (maturation) the Se-enriched yeast in the solution of NaCl, 9 g/L that significantly improved the baking properties. The 100 g of Se-enriched bread contained up to 50 µg Se in the form of SeMet.
ACCEPTED MANUSCRIPT According to Lazo-Vélez, Chávez-Santoscoy, & Serna-Saldivar (2015), the consumption of Seenriched bread could counteract deficiencies - especially in regions that have poor Se soils. However, several factors (i.e., the agronomic, milling, and processing, etc.) significantly affected Se concentration and bioavailability of the flour. Hart et al. (2011) utilized Se biofortified wheat grain cultivated by Broadley et al. (2010) in
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making bread flour for baking Se-enriched bread. A linear correlation was established between the dose of Na2 SeO 4 applied to the crop and the amount of Se in flour and bread indicating a
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minimal loss of Se during grain processing and bread production. SeMet was the predominant Se
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species in the products (flour and bread) obtained from wheat grain biofortified with Se fertilizers. However, SeCys, Se-methylselenocysteine, SeO 3 2-, and SeO 4 2- were other Se species
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detected in the Se-enriched flour and bread. Applying 10 g/ha of Na2 SeO 4 fertilizer raised the
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total Se in wholemeal and white bread by 185 and 155 µg/g, respectively. Supplementing dairy (n = 42) cattle feed with Se-yeast has been shown to increase the amount of
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milk, thus increasing cheese production. The milk Se concentration reached a plateau within a week after commencing Se supplementation. Se-yeast was more effective than inorganic Se
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(Malbe et al., 1995; Ortman & Pehrson, 1999). According to Dillehay et al. (2008), algae have been part of human civilization for thousands of
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years, based on archaeological evidence from 14,000 calendar years before the present (cal yr B.P.) in Chile. China and Indonesia ranked as the top producers of seaweeds and is estimated to value about US $6.7 billion. However, these algae are utilized as foods and food additives (Switzer, 1980; Jassby, 1988; Fournier, Adam, Massabuau, & Garnier-Laplace, 2005; Gantar & Svircev, 2008; Chacón-Lee & González-Mariño, 2010; FAO, 2015; FAO 2016). The demand of algal foods varies across the world due to culture, for instance, typical Japanese diets contain about 9.6 (2014) to 11.0 (2010) g macroalgae per day (MHLW, 2014). Globally the demand for macroalgal and microalgal foods is growing, and they are increasingly being consumed (Wells et al., 2017) or used in formulating functional foods due to its high nutritional value. The schematic
ACCEPTED MANUSCRIPT model of Se metabolism in the algal cell is depicted in figure 3. Detailed Se accumulation and metabolism in algae is reviewed in (Schiavon, Ertani, Parrasia, & Vecchiad, 2017). In aquatic ecosystems, microalgae act as a significant vector of Se accumulation from the watercolumn and partially transform it into organic Se before it is transferred by ingestion to higher organisms. Among the species of microalgae, Spirulina platensis is found to bioaccumulate Se at
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high concentration as 400 mg Se/g dry weight on their cell walls. Scenedesmus quadricauda is reported to also accumulate high Se in the form of Se-enriched microalgal biomass (Fan, Teh,
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Hinton, & Higashi, 2002; Douskova´ et al., 2007; Chen, Zheng, Wong & Yang, 2008; Doucha,
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Lıvansky, Kotrbacek, & Zachleder, 2009).
The European Space Agency, and the National Aeronautics and Space Administration (NASA)
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in the late 1980s and early 90s proposed Spirulina as one of the primary foods to be cultivated
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during long-term space missions due to its nutritional value (Tadros & MacElroy, 1988; Cornet & Dubertret, 1990). Growth media (conditions, i.e., pH, O2 , etc.) can be manipulated to
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significantly increase the nutritional value of algae (Tadros & MacElroy, 1988). Therefore, adding inorganic Se into the growth media could produce Se-enriched algae, which has a great
populations.
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potential of increasing the intake of bioavailable form (organic) of Se by selenium-deficient
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Unicellular marine algae (Dunaliella primolecta, Porphyridium cruentum, and Chlorella vulgaris) have been reported to be excellent accumulators of Se (Bottino et al., 1984; Sun, Zhong, Huang, & Yang, 2014). A significant increase in the cell growth rate and organic Se level of C. vulgaris was reported when exposed to different concentrations of Na2SeO3. Se positively promoted C. vulgaris growth at lower concentrations (≤75 mg L-1), acting as an antioxidant through the inhibition of lipid peroxidation (LPO) and intracellular reactive oxygen species (ROS). The Se-Chlorella could be utilized in designing antioxidative functional foods human health (Sun et al., 2014).
ACCEPTED MANUSCRIPT Spirulina is widely used around the world, because of its rich source of vitamin B12 , in addition to vitamins B1 and B2 . It is estimated to provide one-half of the adult daily requirements of vitamin A (B-carotene) (Tadros & MacElroy, 1988). Arsenyeva & Skripleva (2014) designed yogurt enriched with the supplement «Selenium Algae Plus». A highly bioavailable organic form (SeMet, SeCys) of Se was found to be present in this
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supplement and has a digestibility of 95-98%. The assimilation of Se was found to be inhibited in the presence of sucrose. Therefore, Jerusalem artichoke syrup and stevioside were substituted
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as alternative sweeteners to curb this issue. Nevertheless, these sweeteners can reduce the level
for people who have diabetes and other ailments.
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of sugar in the blood further. The Se-enriched yogurt is not only for mass consumption but also
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The geographical locations, seasons and culture practices (aquaculture) are obstacles in
achieving specific objectives.
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understanding the Se and other nutritional constituents of algae, which impede their utilization in
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POLG mutator mice were used as model organisms during feeding trials to establish the effects of dietary Se status on cardiac gene expression. Diet supplementation significantly (P <0.05, FC
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>1.2) suppressed the expression of age-induced genes that are functionally related to cardiomyocyte apoptosis, hypertrophy, and fibrosis, probably by regulating the transcriptional
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activity mediated by Foxo transcription factors (Xiao et al., 2014). Food processing and storage could potentially alter the nutritive value of algal-formulated foods. Supplementing different forms of Se (Na2 SeO 3 , Se-enriched yeast, lactate protein Se complex, and Se-proteinate) in the feed of pregnant goats revealed elevated Se concentration in the blood of treated animals when compared to the control group. Additionally, kids (about 60% of Se) from treated mothers had higher Se levels in the blood than kids from the control group (Pavlata, Mišurová, Pechová, & Dvořák, 2012). Chlorella sorokiniana is another alga species that can bioaccumulate Se for designing functional food. A Na2 SeO 4 -enriched medium was applied in optimizing growth conditions of C.
ACCEPTED MANUSCRIPT sorokiniana. It was revealed that Se concentration of 100 μg/mL in the medium was the maximum concentration at which growth was maximal. Increasing the concentration above 100 μg/mL caused the collapse (death) of the colonies. Moreover, SeO 3 2- in the medium was biotransformed into mainly SeCys, selenomethyl-selenocysteine (SeMeSeCys), and SeMet, which was confirmed by metallomics analytical approach coupled with HPLC-ICP-MS (Gómez-
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Jacinto, García-Barrera, Garbayo, Vílchez, & Gómez-Ariza, 2012). Microcystins (MCs) in Spirulina are toxic and pose a potential threat to consumers. Roy-
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Lachapelle, Solliec, Bouchard, & Sauvé (2017) detected cyanotoxins in 18 brands of
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Cyanobacteria dietary supplements containing Spirulina and Aphanizomenon flos-aquae. The risks of consuming these products were also assessed. It was discovered that 3 brands out of the
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14 Spirulina products contain MCs ranging between 0.25 and 0.84 µg/g, whereas, 0.8 and 8.2
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µg/g was associated with the A. flos-aquae-based brands. The sum of Spirulina and A. flosaquae-based products MCs varied between 0.01 and 0.63 µg/g and 0.52 and 5.8 µg/g,
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respectively. It was concluded that the quantity of MCs in these brands exceeded the recommended tolerable daily intake (TDI) for adults specified by the World Health Organization
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(exceeding TDI up to 683% for total MCs) (WHO, 2011; Roy-Lachapelle et al., 2017). The amino acid phenylalanine in spirulina causes brain damage - especially to people suffering from
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phenylketonuria (Robb-Nicholson, 2006; Habib, Parvin, Huntington, & Hasan, 2008; Heussner, Mazija, Fastner, & Dietrich, 2012). These toxins could be minimized if organic fertilizers are utilized in cultivating these algae. Chemicals or other additives should be avoided during growing and handling (processing) of algae. There are reports of massive metal accumulation in spirulina supplements and were confirmed in a recent study (approximately 5.1 ppm of the lead (Pb) was detected) (Siva-Kiran, Madhu, & Satyanarayana, 2015). Constructing the cultivation site (pond) far away from industrial settings where the air is less polluted with the residue of heavy metals coupled with using fresh water could be the ultimate remedy for this menace. Nevertheless, using a closed photobioreactor where the environment (internal and external) is
ACCEPTED MANUSCRIPT easily controlled could be an alternative approach. The device could aid in controlling cyanobacterial bloom, therefore minimizing the risk of MCs in the product. The exact doses of MCs are still an open issue for debate, which should be solved in order to prevent possible health risks. Therefore, enacting national and international legislation is necessary to preserve aquatic environments as well as human health (Drobac et al., 2013).
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Se-alga could be useful in designing new functional foods that could exert positive health benefits to consumers. This could be in the form of anti-cancer effects, increase in energy,
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boosting the immune systems, antioxidant proprieties, etc. that will help save huge sums that
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would have ended up in seeking medical care. It could also act as a precursor for selenoprotein synthesis.
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However, in reality, the widespread use of selenized supplements in formulating functional foods
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has a specific limitation. Firstly, the addition of supplements could alter the organoleptic properties of the product. Secondly, some consumers might be allergic (react) to some of these
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products.
According to Rayman (2004), the initial production of selenized supplements was not meant for
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only human consumption. However, the increase in demand for these products across the world following the approval from the US Food and Drug Administration has shifted perception on
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these supplements. The leading manufacturers of these supplements are Pharma Nord, Vejle, Denmark, Alltech, Lexington, KY, USA, Lallemand, Montreal, Canada, etc. Manufacturers are cautioned to follow international standards in order to avoid contaminations, i.e. microbial or heavy metals.
The tolerable upper intake level (UL) of Se for adults in the US is 400 µg (5.1 µmol)/day based on selenosis as the adverse effect. UL for infants of the age 0–6 and 7–12 months are 45 µg (0.57 µmol)/day and 60 µg (0.76 µmol)/day of selenium, respectively. Children of the age 1–3, 4–8, 9– 13 years has UL of 90 µg (1.1 µmol)/day, 150 µg (1.9 µmol)/day, and 280 µg (3.6 µmol)/day of selenium, respectively (Food and Nutrition Board Institute of Medicine, 2000).
ACCEPTED MANUSCRIPT The European Commission health and consumer protection directorate have derived the UL of 300 µg Se/day for adults, pregnant and lactating women based on no-observed-adverse-effect level (NOAEL) of 850 µg/day for clinical selenosis in the study on 349 subjects of Yang et al. (1989b) and Yang & Zhou (1994). In addition, UL of 250, 200, 130, 90, 60 μg selenium/day was derived for Europeans of the age 15-17, 11-14, 7-10, 4-6 and 1-3 years respectively (European
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Commission Health & Consumer Protection Directorate, 2000; European Food Safety Authority, 2006). Some European Union countries have set the mean daily intakes of Se for non-vegetarian
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adults as: United Kingdom 63 μg/day, France 29-43 μg/day, Sweden 24-35 μg/day, Belgium 28-
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61 μg/day, Denmark 41-57 μg/day, Finland 100-110 μg/day, The Netherlands 40-54 μg/day, Norway 28-89 μg/day, and Spain 79 μg/day (Alexander & Meltzer, 1995; van Dokkum, 1995;
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Johansson et al., 1997). The Se fertilization programme in Finland appeared to have successfully
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raised Se status in the population (100-110 μg Se/day). However, in the case of Norway, this could be due to the continuous importation of biofortified wheat (European Commission Health
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& Consumer Protection Directorate, 2000; Stoffaneller, & Morse, 2015). The recommended UL by the Americans (400 µg) is higher when compared to the EU value (of 300 µg Se/day).
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3.1. Preparation of selenized Tom-brown porridge flour Tom-brown flour is a powder made by grinding roasted raw grains and is used for making
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different foods. This flour is the main ingredient for porridge which nursing mothers used in weaning their babies in Africa. Nursing mothers could also use this porridge in supplementing the breast milk for their babies since it expensive buying products like milk (NAN milk), cerelac, etc., which are formulated in other parts of the world. Figure 4 shows the entire process of producing the Tom-brown porridge flour. 1. After harvesting and drying of cereals (yellow corn and guinea corn). Screening is followed to remove stones and other foreign particles. Further cleaning such as winnowing is carried out. The cereal is then roasted in a frying pan on a fire until it turns golden brown. It is then removed and allowed to cool.
ACCEPTED MANUSCRIPT 2. Depending on the location, legumes like groundnuts, soybeans, etc. could be added in the mixture. Similar treatments as explained above are replicated here. 3. The roasted cereals and the legumes are mixed and milled into flour using either an electric or a diesel-milling machine. Some producers do pound the grains in mortar and pestle to separate the chaff. However, they are criticized of eliminating vitals nutrients
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that might have attached to these chaffs. 4. Pasteurized dried selenized supplements (yeast and algae) are then mixed with the milled
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flour (Selenized enriched Tom-brown).
4. Extraction and quantification of organic Se.
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5. The selenized enriched Tom-brown flour is now ready for consumption.
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Several researchers have outlined methods for quantifying Se in various matrixes (Uden,
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Boakye, Kahakachchi, & Tyson, 2004; Kaplan, Gil, & Cerutti, 2006; Harwood & Su, 1997; Goenaga-Infante et al., 2008; Pedrero, & Madrid, 2009; Soruraddin, Heydari, Puladvand, &
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Zahedi, 2011; Chen et al., 2011; Nam, & Basu, 2011; Duan, He, & Hu, 2012; Fagan et al., 2015; Jagtap & Maher, 2016; Tie et al., 2016). The chemical form of Se in sample, matrix (i.e., plant,
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fish, meat, yeast, egg, human serum, blood, etc.), the instrumentation availability, experience of the researcher are factors to consider when selecting methods for sample treatment for the
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isolation and identification of the Se species (Pedrero & Madrid, 2009). According to Peachey, McCarthy,
& Goenaga-Infante, (2008) selecting extraction methods, which exhibit high
extraction efficiency while preserving the integrity of Se species is essential for the accurate measurement of Se species. Extraction of Se species using aqueous solutions (water, water– MeOH, and neutral buffers), acid–alkali hydrolysis (HCl, methanesulfonic acid, tetra-methylammonium hydroxide (TMAH)), and enzymatic hydrolysis (proteinase K, subtilisin, pepsin and protease, or mixture of enzymes protease and lipase) are the methods available currently. Acidic extraction is characterized with low SeMet recoveries, in some instance complete loss of SeCys (~100%) with extracts containing elementary Se0 (Huerta, Sanchez, & Sanz-Medel, 2004; Yang,
ACCEPTED MANUSCRIPT Sturgeon,
McSheehy,
& Mester,
(Selenomethyl-selenocysteine,
2004; Jagtap
& Maher,
γ-glutamylselenomethyl-selenocysteine,
2016). Selenoamino acids etc.)
are
water-soluble
hence, are aqueously extracted with hot water with excellent recovery. However, excellent recovery is only achieved if Se species are not incorporated into proteins (Pedrero & Madrid, 2009). Therefore, coupling aqueous with enzymatic treatment could improve the extraction
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efficiency. Enzymes can interact, creating pores on proteins housing Se species, thereby resulting in leaching. Organic Se was successfully extracted from selenized Yeast (McSheehy, Szpunar,
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Haldys, & Tortajada, 2002; Encinar et al., 2003), plants (Zhang & Frankenberger, 2001) and
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mushrooms (Stefanka, Ipolyi, Dernovics, & Fodor, 2001) using an aqueous method with good recoveries. Previous findings (Moreno, Quijano, Gutiérrez, Pérez-Conde, & Cámara, 2004;
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Huerta, Sanchez, & Sanz-Medel, 2004; Dumont, Vanhaecke, & Cornelis, 2006) have suggested
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that the incorporation of sodium dodecyl sulfate (SDS) increased/improved the extraction efficiency up to 50%. SDS interact with proteins thereby rendering them soluble in water hence
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increasing the release of selenoproteins. Chen et al. (2011) modified the method previously described by Dumont et al. (2005) to isolate Se species from both yeast and clover. High purity
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deionized water (5mL) was added to a 25 mg sample in a centrifuge tube, followed by a hot water bath at 50 ◦C for 24 h. The extract was then centrifuged and filtrated, and the filtrations
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were made up to 10mL with deionized water. The enzymatic hydrolysis is the preferred method of many researchers because it presents several merits compared to the other methods. Enzymatic hydrolysis is selective and specific because enzymes act only on specific chemical bonds and can therefore distinguish between fractions of elements associated with the different components of the sample matrix. Furthermore, elemental losses by volatilization are minimized because enzymes perform well at moderate pH and temperature.
Reagents are not required to neutralize the excess of acid or
alkali in acidic extracts before chromatography analysis hence reducing any risk
of
ACCEPTED MANUSCRIPT contamination (Bermejo-Barrera, Fernandez-Nocelo, Moreda-Pineiro, & Bermejo-Barrera, 1999; Jagtap & Maher, 2016). The Se yeast sample could accurately be measured (0.04 g) into microcentrifuge tubes. Protease enzyme solution (13.33 mg Protease XIV in 0.5 mL of Tris buffer pH 7.5) is then added and vortex mixed for 2 min. Samples were ultrasonicated for 25 s at 80 % amplitude, a power output
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of 30 W for 15 minutes. Ice cubes is used to cool the sample from the heat generated due to the ultrasonication. Before commencing the sonication, 250 μL enzyme solution is used to rinse the
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probe. Extracted samples is then centrifuged at 14,000 rpm for 3 min. Supernatants are then
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transferred to 15 mL centrifuge tubes. Pellets washed with water (0.9 mL) and vortex-mixed completely in solution. This step is repeated thrice. Volumes are adjusted to 15 mL using water
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and mixed thoroughly before removing an aliquot (2 mL) for filtration (0.25 μm) and dilution
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before analysis (Fagan et al., 2015). Residual yeast, white wine, and beer were subjected to enzymatic hydrolysis before quantification and identification of selenium species (Pérez-Corona
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et al., 2011; Sánchez-Martínez et al., 2012). The extracts are then injected into high-performance liquid chromatography with inductively coupled plasma mass spectrometry (HPLC-ICP-MS)
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using a sample injection valve fitted with a 100 µL loop. Details of the separation columns and operating conditions are reported in Pérez-Corona et al. (2011) and Sánchez-Martínez et al.
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(2012). Comparing the retention time and spiking experiments Se species are identified. Se species (Se(VI) and SeMet from impurities in the protease hydrolysis is used as blank for the analysis. High-resolution Time-of-flight Mass Spectrometry could also be used in quantifying organic Se (Krittaphol et al., 2008). Good Se recoveries by enzymatic hydrolysis of yeast, fish and plant CRMs and miscellaneous have been reported. Approximately 96% and 93% of total Se in Antarctic krill and cod muscle (CRM BCR 422) were documented, respectively (Siwek, Galunsky, & Niemeyer, 2005; Kapolna, Gergely, Dernovics, Illes, & Fodor, 2007). Hydride generation–inductively coupled plasma optical emission spectrometry (HG-ICPOES) was proposed as an effective method of
ACCEPTED MANUSCRIPT determining total Se in the sample. This system allows the determination of Se in high copper dose samples. High precision and accuracy were obtained after analyzing reference samples (Kaplan, Gil, & Cerutti, 2006). The outlined approaches provide a secure, rapid, reproducible, and cost-effective platform for computing organic Se. However, acidic hydrolysis of Se-enriched foods was found to be the most effective technique of measuring total Se. Therefore, coupling
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two pretreatment methods are recommended for effective extraction of Se species from samples. 5. Se bioavailability
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Ruby et al. (1999), defined bioavailability as the fraction of a substance released from food
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matrix that is absorbed via the intestinal barrier (bioavailable fraction), transported into the bloodstream or an organ, and is available to promote its activities in the exposed organism (Ruby
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et al., 1999). However, the non-accessible fraction is not absorbed, thus is directly excreted
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(Moreda-Pineiro, Moreda-Pineiro, & Bermejo-Barrera, 2017). The fraction of a nutrient or substance in food that is theoretically soluble from its matrix and can quickly enter the
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bloodstream or affect the gastrointestinal (GI) tract is termed bioaccessibility (Ruby et al., 1999; Oomen et al., 2002; Stahl et al., 2002; Moreda-Pineiro et al., 2017). The physiological state of
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the consumer (i.e., age, health status, gender, etc.,), GI conditions, the form of Se in food (Ziegler et al., 1978), and geographical location are factors affecting the bioavailability of any
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food.
With the aid of isotopically or non-isotopically labeled Se (Miller et al., 1981; Ruby et al., 1999; Fernandez-Garcıa et al., 2009) several stimulated studies have been carried out to assess the bioavailability of Se in natural products or Se-enriched, raw and processed products. Recently Moreda-Pineiro et al. (2017) published a good review on Se bioavailability in various food products. In vivo, human tests show a significant increase in the Se levels in plasma or whole blood after supplementation
with
Se-fortified
wheat
and
process-fortified
Se
biscuits.
Plasma
Se
concentrations with Se-fortified wheat was more than (from 122 to 194 µg/L) with process-
ACCEPTED MANUSCRIPT fortified Se biscuits (from 122 to 140 µg/L). This could be as a result of different species of Se available in the products (i.e., SeMet and SeOMet in Se-fortified wheat and process-fortified Se biscuits, respectively) (Thompson et al., 1985; Mutanen, 1986; Van der Torre et al., 1991; Djujic et al., 2000; Kirby et al., 2008). A similar study was carried out on rats using tuna and Sefortified wheat. There was a significant decrease in glutathione peroxidase (GSHPx) activity
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with groups treated with Se-fortified wheat compared with the tuna treated groups, suggesting that Se in wheat was more available than that in tuna (Alexander, Whanger, & Miller, 1986).
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Yoshida and colleague (1999) quantified tissue Se accumulation and GSHPx activity of 4-week-
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old male Wistar rats fed with high-Se yeast (SeY) and Na2 SeO 3 . Strong correlations were established between the supplementary Se levels and the tissue Se contents or GSHPX activities,
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indicating that that Se in SeY is more bioavailable than Na2 SeO 3 . Therefore, SeY should be used
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in formulating Se functional food.
Selenium-deficient rats showed dose-dependent increases in GSHPx activities in blood and in
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thioredoxin reductase activity in the liver after feeding them with yellow peas and oats harvested from the high-Se soil of South Dakota, the United States. It was concluded that Se from yellow
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peas and oats was highly bioavailable (Yan & Johnson, 2011). In a similar study, soy protein isolates (from soybean) was more Se bioavailable relative to tofu (Yan, Reeves, & Johnson,
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2010).
Bioavailability of Se (organic Se species such as MeSeCys, SeCys2, and SeMet) in Allium was found to be very high after an in vitro gastric and gastrointestinal digestion (proteolysis) than in Chive. Speciation studies revealed that these Se fortified Allium was not safe for human consumption because Se isolated was in the form of selenite. However, 90% of the total Se content of both samples became bioaccessible (Kapolna & Fodor, 2007). This finding was in agreement with previous studies (de Leon, Sutton, Caruso, & Uden, 2000; Kotrebai, Birringer, Tyson, Block, & Uden, 2000; Shah, Kannamkumarath, Wuilloud, Wuilloud, Caruso, 2004; Wrobel et al. 2004).
ACCEPTED MANUSCRIPT Selenized yeast and yeast-based nutritional supplements have been extensively assessed using in vitro approaches. Supplementation of the human diet with yeast and selenized tablets revealed that Se levels in plasma and whole blood increased significantly, indicating high Se availability in yeast (Alfihan, Aro, Arvilommi, & Huttunen, 1991; Xia, Zhao, Zhao, Zhu, & Whanger, 1992; Thompson, Robinson, Butler, & Whanger, 1993; Alfthan et al., 2000; Larsen et al., 2004; Bugel
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et al., 2008; Bost & Blouin, 2009; Mahmoud, 2012). In vivo animal (pig, ewe, rat, catfish, yellowtail kingfish, trout, and boiler chicken) studies showed that Se in yeast was highly
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bioavailable. Moreover, Se in yeast was reported to be almost twice as bioavailable as Se from
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Na2 SeO 3 and Na2 SeO 4 . High bioaccessible percentage (55–89%) was reported in the case of total Se in yeast, whereas low bioaccessible percentages (26–41%) were attributed to SeMet in yeast
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and yeast-enriched supplements (Smith & Picciano, 1987; Wang & Lovell, 1997; Yoshida et al.,
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1999; Dumont, Vanhaecke, & Cornelis, 2004; Hinojosa-Reyes, Ruiz-Encinar, Marchante-Gayon, Garcıa- Alonso, & Sanz-Medel, 2006a; Hinojosa-Reyes, Marchante-Gayon, Garcıa-Alonso, &
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Sanz-Medel, 2006b; Mateo, Spallholz, Elder, Yoon, & Kim, 2007; Wang & Xu, 2008; Rider et al., 2009; Rider, Davies, Jha, Clough, & Sweetman, 2010; Jang et al., 2010; Hall et al., 2012;
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Briens, Mercier, Rouffineau, Vacchina, & Geraert, 2013; Le & Fotedar, 2014). Hence, selenized yeast could be exploited to produce Se fortified functional food, which can serve as a dietary
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source of Se intake.
Food processing (i.e., cooking, frying, grilling, milling, fermentation, etc.) could affect the bioavailability of Se (Moreda-Piñeiro et al., 2017). Cooked tuna had slightly higher Se bioaccessible than in the uncooked sample. The heat from cooking might have affected the efficiency of digestive enzymes, hence improving protein digestibility, which facilitated Se release to enhance the bioaccessibility (Cabanero et al., 2004). In vivo, Se-deficient male albino rats showed higher bioavailability of Se in raw salmon than cured salmon. Curing salmon might have altered some enzymes, which could have inhibit the release of Se. However, Se from raw and cured salmon tended to be more bioavailable compared with Na 2 SeO 3 . The induction of
ACCEPTED MANUSCRIPT plasma GSHpx activity and Se accumulation in the femur and muscle goes to support the results (Ørnsrud & Lorentzen, 2002). Shi & Spallholz (1994) observed a significant decrease in liver GSHPx activity when Fischer 344 rats were fed a Se-deficient diet than the control rats. Upon supplementing the diets with 0.10 mg/kg Se as Na2 SeO 3 , Na2 SeO 4 , raw or cooked ground beef, liver GSHPx activity began to
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increase on treated rats than on the control rats. The percentage of liver GSHPx activity recovery were as follows: cooked ground beef (139%, p < 0.05), raw beef (127%, p < 0.05), Na2 SeO 4
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(117%, p < 0.05), and Na2 SeO 3 (98%, p > 0.05) indicating the bioavailability of Se from cooked
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ground beef and raw beef were greater than Na2 SeO 3 or Na2 SeO 4 . The cooking might have activated certain enzymes to hydrolyze all the peptide bonds in proteins, magnifying Se release
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thus increase its bioaccessibility. Milling has been shown to decrease the amount of Se in cereal
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product. The losses appear not to be excessive since Se is evenly distributed throughout the wheat kernel. Nevertheless, the Se that was lost to the consumer due to milling appeared in the
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by-products used as animal feeds, therefore some of this Se would eventually be in the meat and ultimately utilized by the consumer (Ferritti & Levander, 1974). According to Pederson et al.
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(1991) and Fairwetaher-Tait (1997), the presence of arsenic, vitamin C and guar gum (guaran) in the diet inhibit the uptake and assimilation of Se making it non-bioaccessible to the consumer.
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6. Mechanistic understanding of Se toxicity and health concern It is scientifically proven that organic Se (Se-yeast) is less (not toxic) than inorganic Se. Humans and animals (i.e., farm animals and poultry) are susceptible to excess or insufficient Se in the diet. High intake of organic and inorganic forms of Se have similar effects (Institute of Medicine, Food and Nutrition Board, 2000). The toxicity of Se depends on many factors, and it is difficult to make categorical pronouncements about the quantities of Se involved. The quantity and chemical forms of Se consumed coupled with other elements i.e. arsenic mercury, cadmium, etc. and dietary constituents could determine the class under which the toxicity (acute and chronic toxicities) is categorized (Diplock & Hoekstra, 1976; Farzaneh, 2015).
ACCEPTED MANUSCRIPT Seafood and meats (organ) are rich sources of Se in food. Other sources include muscle meats, grains, and dairy products (Table 2). However, care should be taken when choosing these products since some contain an excess amount of Se. For example, Brazil nuts contain around 68–91 mcg per nut and could lead to Se toxicity (poisoning) if consumed on a regular basis. In a study, when 201 people consumed a liquid dietary supplement containing 200 times Se than the
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recommended level, they experienced severe adverse reactions. i.e., vomiting etc. (Sunde, 2006; Sunde, 2010; MacFarquhar et al., 2010; Sunde, 2012).
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Saha, Fayiga, Hancock, & Sonon, (2016) discovered that the minimum lethal dose of Se in the
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form of Na2 SeO 3 varied from 1.5 to 8.0 mg/kg body weight in farm animals. Acute toxicity studies showed the LD50 for Se-yeast was 37·3 mg/kg compared with 12·7 mg/kg for Na2 SeO 3
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(toxic) (Figure 5). This evidence proved that Se-yeast is considerably less toxic than Na2 SeO 3
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(Vinson & Bose, 1987; Spallholz & Raftery, 1987).
When Se is consumed, be it inorganic (selenite) or organic (selenomethionine, selenized-yeast
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cells, etc.), it is metabolized by reduced glutathione (GSH) to yield seleno-diglutathione, which is then transformed into seleno persulfide (GSSeH). Under anaerobic conditions, GSSeH could
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enzymatically convert into volatile hydrogen selenide (H2 Se) or decays spontaneously to Se and GSH. Na2 SeO 4 is assimilated by active transport in the intestinal brush border whereas via
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passive diffusion, Na2 SeO 3 is absorbed. H2 Se enters the host cell with efficiency and exacerbates oxidation of GSH (Ganther, 1971; Hsieh & Ganther, 1975; Tarze et al., 2006). According to Shelly (2009), GSH is a ubiquitous intracellular protein with diverse functions that include detoxification,
antioxidant defense,
maintenance of thiol levels,
and modulation of cell
proliferation, etc. Se can combine with GSH and other thiols to form glutathioselenol, which could further reduce GSH into H2 Se. ROS such as superoxide ions (O 2 •), hydrogen peroxide (H2 O2 ), and hydroxyl radicals (OH•), are likely to be generated from the reaction between H2 Se and molecular cell oxygen. The reaction generates oxygen for the continuous oxygenation of glutathione resulting in the depletion of intracellular GSH, S-adenosylmethionine, which inhibits
ACCEPTED MANUSCRIPT protein synthesis, resulting in the complete substitution of sulfur (trans-sulfuration pathway) by Se in cellular metabolism. This leads to the loss of enzymatic activity in some sulfur-containing proteins due to the replacement of sulfur in sulfhydryl groups or thiols (vital for the formation of a disulfide bond) with Se. The consequence is genotoxic/cytotoxic effects due to oxidative stress on the cells (Vernie et al., 1974; Stadtman, 1974; Hoffman, 1977; Anjaria & Madhvanath, 1988;
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Seko, Saito, Kitahara & Imura, 1989; Stadtman, 1990; Yan & Spallholz, 1993; Spallholz, 1994; Seko & Imura, 1997; Pinson, Sagot & Daignan-Fornier, 2000; Tarze et al., 2006).
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Toxicity may also arise due to the reactions of H2 Se with metal-containing proteins, and this was
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confirmed by in vitro experiments. Recent studies showed that the iron atom in lipoxygenases of human monoclonal B-lymphocytes was sensitive to H2 Se. It was also reported that H2 Se
consequently affecting the human hepatoma cells (Eghbal,
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altering the redox reaction,
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exacerbates the inhibition of heme-containing enzymes, which belong to the respiratory chain,
Pennefather & O’Brien, 2004; Bjornstedt et al., 1996). These reactions could lead to dizziness,
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unconsciousness, etc. Pancreatic ribonuclease A was almost completely inactivated when treated with selenious acid (Ganther & Corcoran, 1969). Se toxicity could result in cell damages of
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genetic information (genotoxicity), which causes mutations. The availability of excess ROS in the cell reacts with both deoxyribose and bases in DNA, causing base lesions and strand breaks.
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Nevertheless, ROS can significantly oxidize DNA, thus inhibiting DNA repair, gene regulation (down-regulated) and threatening gene stability (Ramana, Boldogh, Izumi, & Mitra, 1998; Kelly et al., 2008; Wei et al., 2001; Zhou et al., 2003). In-vivo and in-vitro studies have confirmed that liver glutathione (thiols) producing organ is susceptible to Se toxicity due to its involvement in the rapid catalytic reaction of Se compounds. Thiol-containing enzymes like methionineadensyltransferase,
succinate-dehydrogenase,
NADP+-isocitrate-dehydrogenase,
and
lactate-
dehydrogenase were altered via Se toxicity (Nebbia et al., 1990; Mezes & Balogh, 2009). According to the literature, the minimum toxic dose of Na2 SeO 3 can cause clinical changes in birds, mg/kg body weight: ducklings – 9.4; broilers – 1.7; laying hens – 33.4; indices – 0.9. The
ACCEPTED MANUSCRIPT oral LD50 Se for young turkeys, ducks, chickens are 13.5, 64.0, and 9.7 mg/kg live weight, respectively (Surai, 2002; Sobolev et al., 2018). Feeding laboratory rats with SeO 3 2- and Se-yeast for 8 weeks manifested severe cardiotoxicity, hepatotoxicity, and splenomegaly in rats fed with SeO 3 2- whereas no such symptoms were exhibited in rats fed with the latter. Further histological examinations revealed that Se-yeast had
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greater deposition of Se. (Spallholz & Raftery, 1987). Clinical symptoms of Se poisoning in farm animals (cattle and sheep) are excessive salivation,
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garlic smell when exhaled, vomiting, loss of appetite, difficulty in breathing (shortness of
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breath), seizures, hoof deformation, lameness, diarrhea, alopecia (hair loss), bloating, loss of vision, exhaustion, paralysis and death, often from respiratory failure (Sobolev et al., 2018).
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Functions and tertiary enzymes structure are altered due to the formation of Se trisulfide
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complex (Se + Cys + coenzyme A). This could compromise the immune system due to the inability of the enzymes to catalyze activities, i.e., digestion, breathing, nutrient, absorption, etc.
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Moreover, Se toxicosis can lead to the production of methyl selenide (Se2-), a precursor for inducing oxidative stress on the system (Mézes & Balogh, 2009; Sobolev & Pacelja, 2016).
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A controlled clinical trial of testing a chemopreventive agent against prostate cancer progression by supplementing the diet of 24 men with either 1600 or 3200 µg/day of Se-yeast was carried
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out. The group fed with 3200 µg/day were reported to show symptoms such as garlic breath, brittle nails, and hair, stomach upset and dizziness unlike the former group (Reid et al., 2004). No side effects were documented when a similar study was replicated on hepatitis-B surfaceantigen positive patients (n=226) (Yu, Zhu, & Li, 1997). The latter researchers utilized 200 mg/d Se whereas the former supplemented with 3200/1600 mg/d. Other factors such as racial difference, age, health status, etc., could all be attributed to these findings. Rigorous studies are needed to elucidate the toxicity levels of Se taking into consideration the factors above (racial difference, age, health status). Table 3 shows the recommended daily Se intakes in some selected countries.
ACCEPTED MANUSCRIPT 7. Impact Se on gut microbiota. Microorganisms constitute the gut microbiota of every mammal and are essential in digestion, recycling of nutrients, and inhibiting colonization by pathogenic organisms (Guarner and Malagelada, 2003). Microbes residing in the gut are known to be actively involved in drug, Se, and vitamin metabolism (Boyle et al., 2006; Jia et al., 2008; Kasaikina et al., 2011b). Previous
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works have shown that Se administered as SeMet and other forms undergo a significant enterohepatic recycling, which involved the gut microflora (Xia et al., 1992; Krittaphol et al.,
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2009; Krittaphol, McDowell, Thomson, Mikov, & Fawcett, 2011a; Kasaikina et al., 2011; Saini
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& Tomar, 2017).
It was reported that Se could alter the composition of the gut microbiota, consequently
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influencing the selenoproteome and Se levels of the host. However, the composition of the diet
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coupled with other trace elements could influence the bacterial population in the gut (Kasaikina et al., 2011). Therefore, Se could impact commensal bacteria in the host that regulate nuclear
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factor-kB (NF- k B) and peroxisome-proliferator-activated receptor (PPARγ) which aids in the activation of immune cells during inflammatory bowel disease (IBD) (Nettleford & Prabhu,
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2018).
Gut microbiota contributes to the excretion of excess Se through the production of methyl Se
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compounds and elemental Se, thus protecting the host from selenosis (Krittaphol, McDowell, Thomson, Mikov, & Fawcett, 2011a). Probiotics like LAB strains improve the degradation of proteins in milk products, thus increasing the bioaccessibility of Se (Shen et al., 1996; Saini & Tomar, 2017). Enterococcus durans LAB18s was proposed as a potential probiotic that could be used for designing Se-enriched products. The highest percentage of bioaccessible organic Se was found in the fraction of total protein, followed by the fraction of polysaccharides and nucleic acids (Pieniz et al., 2017). The microbiota diversity could increase, thereby keeping the integrity of the gut microbiota intact due of Se bioavailability. Disruption of intestinal microbiota could lead to diseased states and
ACCEPTED MANUSCRIPT disorders (Vijay-Kumar et al., 2010; Kasaikina et al., 2011; Virgin & Todd, 2011). Motor control and anxiety behavior were reportedly correlated to host microbiota (Diaz Heijtz et al., 2011) In vitro probiotic tests confirms the ability of LAB to survive in the simulated gastric juice at pH 2 and could be exploited for the development of Se enriched functional foods (Se-enriched yogurt, kefir, beer, etc.) as a source of dietary Se (Saini, Tomar, Sangwan, & Bhushan, 2014; et al., 2014; Saini & Tomar, 2017). According to Saini et al. (2014), L. reuteri
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Pophaly
NCDC77 is a novel strain with greater potential to biotransform inorganic Se from a medium
(CH3 SeCys)
and
the
latter
transforms
to
highly
bioaccessible
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methylselenocysteine
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than the control bacteria. Additionally, it synthesizes selenocysteine (dimer of SeCys), and
methylselenol (efficient than SeMet) by the action of β-lyase and is considered as the most
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effective form of organic Se utilize in cancer therapy.
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In vitro and in vivo studies demonstrate that a variety of probiotic bacteria can metabolize SeMet and Na2 SeO 3 to produce methylated Se compounds. Probiotic mixtures significantly increased
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Se content in the liver and decreased them in the kidney after administration of SeMet compared to the control rats (Krittaphol et al., 2011b). In a similar study, an improved HPLC method was
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proposed for the quantification of L-SeMet in rat gut content suspensions prepared from the different parts of the intestine (jejunum, ileum, caecum, and colon). L-SeMet metabolism in the
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gut occurs mainly in the caecum because it houses microorganisms with high metabolic activity towards L-SeMet. Moreover, caecum is known to play a vital role in digestion and therefore contains a myriad of enzymes produced/secreted by the gut microbiota (Hall, Youngs, Keighley, 1992; Kurosawa, Ikeda, Sukemori, & Kurihara, 2005; Krittaphol et al., 2009). Further work is necessary to clarify into the biotransformation of L-SeMet by gut microflora (Krittaphol et al., 2009) about the bioavailability of Se. Lastly, in vivo and in vitro elucidation of Se metabolism pathway of LAB strains to get molecular and technological insights on prospective cultures and to model their response in fermented foods (Pophaly et al., 2014) need to be considered in future.
ACCEPTED MANUSCRIPT 8. Factors to consider when designing functional food 1. The enrichment/fortification must not alter the organoleptic properties (flavor, color, texture or odor) of the original food. 2. The active ingredient (Se) should demonstrate acceptable stabilities in the fortified foods. 3. The potential for any interactions that could occur between micronutrients in multiple
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fortified systems, as well as interactions between the active ingredient and food vehicle that may interfere with the metabolic uptake of the active ingredient, also need to be
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thoroughly examined.
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4. The addition of extra additives such as binders encapsulates, and stabilizers to improve the retention of the active ingredient should not require significant changes to existing
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technologies.
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5. The functional food must be well absorbed from the food vehicle at the level of consumption compatible with a healthy diet.
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6. The final cost of designing functional foods must not decrease the affordability of the food or increase the competitiveness with the unfortified alternative (Sangakkara, 2011).
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9. Conclusions and future trends
In developed nations, functional foods are considered to be and are ranked as one of the critical
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state policies. Adequate nutrition promotes growth and development, prevents diseases and increases productivity and life expectancy. Health risks associated with Se deficiency have affected many lives negatively and remains a threat to humanity. Therefore, enriching foods, which are consumed on a regular basis by a large portion of the population with Se, is the ultimate solution in combating the menace of Se deficiency.
Brewing yeast has the necessary
enzymes to biotransform inorganic Se (Na2 SeO 3 ) to organic form which is incorporated in the aged beverage. Se-enriched beverages are safe for consumption; hence, consumers should not worry about the issue with toxicity. S. cerevisiae was found to possess better biotransformation efficiency than the other yeast species. Selenized yeast and algae could be taken as supplements
ACCEPTED MANUSCRIPT or as food additives. However, the safety of Se-algae is not guaranteed because, recently, there have been some reports about MCs in Spirulina. Consumers should be careful about the source of Se-algae they are purchasing since the concentration of MCs depends on several factors, i.e. geographical location, strain, type of fertilization practice, etc. The public is again advised against Brazil nuts. It contains about 544 mcg of Se which can cause Se poisoning.
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Recent advances in the field of nutrigenomics are vital tools for investigating functional food. Deciphering the interactions between bioactive compounds (Se in this study) and proliferation of
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diseases based on an individual’s genetic profile is the ideal strategy to maximize health benefits
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and reduce disease risk from this food. The application of Nutrigenomics in the study of antiaging, obesity, cancer, etc. coupled with Se-enriched (functional food) is the best tool to
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understand the comprehensive benefits and side effects an individual gets from these foods.
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Thanks to Ventor and colleagues who completed the human genome sequence in 2001 (The Celera Genomics Sequencing Team, 2001), hence making the study of Se on individual genetics
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feasible.
The introduction of systems biology coupled with biotechnology is another vital tool in the
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future advancement of functional foods. Metabolic pathways could be engineered either to suppress the expression of genes responsible for the accumulation of excess Se in crops (i.e.,
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Brazil nuts) or to develop new strains of probiotics capable of secreting organic Se in the products. Nevertheless, this tool could also be used in developing genetically modified produce enriched with Se and other vital minerals, which could potentially improve the health of the consumer.
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Authors’ Contributions
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AP, NVD, KYM, and EFK have contributed to the study conception and design; AP searched
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for literature and wrote the manuscript; NVD, KYM, and EFK were responsible for the graphs, tables, and references. All authors have read and approved the final manuscript.
Acknowledgments
This work was financially supported by the Government of Russian Federation, Grant 08-08.
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Tables: Table 1. Total selenium in selenized beverages Se (IV) added
Se in selenized
(µg/mL)
beverages (µg/mL)
0.1
0.086±0.003a
1.0
0.61±0.08a
2.0
1.1±0.4a
10.0
6.0±0.7a
20.0
6.0±0.5a
30±5
50
15±1b 30±6b
30.6
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200 300
55±4 60±7
47±5b
30.8
115±8b
57.5
167±6b
55.6
300±10b
60.3
300±10b
29.7
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500
61±3
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43±5
30.3
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100
1000
% Se
Modified with permission from ref 4390651395301(Elsevier, 2011) and 4390660155828 (Elsevier, 2012). a
selenized-enriched beer after 12 days fermentation (Sánchez-Martínez et al., 2012)
b
selenized-enriched wine after 21 days fermentation (Pérez-Corona et al., 2011)
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Table 2. Different sources of functional foods/beverages containing selenium Reference
Se-enriched mushroom
Ogra, Ishiwata, Encinar, Lobinski, & Suzuki, 2004
Se-fortified yogurt
Achanta, Aryana, & Boeneke, 2007
Se-enriched eggs
Fisinin, 2007; Gajčević, Kralik, Has-Schön, & Pavić,
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2009; Fisinin, Papazyan, & Surai, 2009; Bennett &
Hu, Xu, & Pan, 2001; Xu et al., 2003; Vodnar &
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Se-enriched tea
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Cheng, 2010
Socaciu, 2014
Lintschinger, Fuchs, Moser, Kuehnelt, & Goessler,
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2000 Se-biofortified apples
Wortmann & Daum, 2018
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Se-biofortified prickly pear
Tyrala, Borschel, & Jacobs, 1996
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Se-fortified infant formula Se-enriched fermented milk
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Se-enriched sprout
Alzate, Perez-Conde, Gutierrez, & Camara, 2010 Banuelos et al., 2011 IP & Lisk, 1993; Ip & Lisk, 1995, Lu et al., 1996
Se-biofortified onion
Adhikari, 2012; Golubkina et al., 2016
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Se-enriched garlic (Se-garlic)
Se-biofortified broccoli
Pedrero, Elvira, Camára, & Madrid, 2007; Adhikari, 2012
Columbus Egg
De Meester et al., 2000
Se-enriched beef
Lawler, Taylor, Finley, & Caton, 2004; Lee et al., 2006; Taylor et al., 2008
Se-enriched wine
Pérez-Corona et al., 2011
Se-enriched beer
Sánchez-Martínez et al., 2012
Se-enriched kvass
Muravyov, 2018
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Liangcai, 2015; Wave et al., 2017
Se-enriched bread
Sarudi, Lassu-Merényi, Rumi, & Kelemen, 1997; Bryszewska et al., 2007; Hart et al., 2011; Lazo‐ Vélez, Chávez‐ Santoscoy, Serna‐ Saldivar, 2014 Yan & Johnson, 2011
Se-biofortified shallot
Ogra, Ishiwata, Iwashita, & Suzuki, 2005
Se-biofortified pumpkin
Smrekolj, Stibilj, Kreft, & Kápolna, 2005
Se-biofortified dill
Cankur, Yathavakilla, & Caruso, 2006
Se-biofortified chicory
Mazej, Falnoga, Veber, & Stibilj, 2006
Se-biofortified radish
Ogra et al., 2007
Se-biofortified ramp
Whanger, Ip, Polan, Uden, & Welbaum, 2000
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Se-fortified oat
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Table 3. Current recommended selenium intakes for adults (µg/d) Male
Females
USA and Canada, 2000
55
55
Australia, 1990
85
70
Belgium, 2000
70
70
30-70
30-70
55
55
60
50
55
55
55-60
45
65
55
50
40
75
60
40
30
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Countries
Germany, Austria, Switzerland, 2000
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EC Scientific Committee on Food, 2003
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France, 1996 Italy, 1999
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Japan, 1999
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New Zealand and Australia (proposed levels) Nordic countries, 1996
World
Health
Organization/Food
and
Agriculture
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Organization/International
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UK (Committee on Medical Aspects of Food Policy, 1991
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Atomic Energy Agency, 1996
Adapted with permission from ref 4392370266741 (Cambridge University Press, 2006) (EC Scientific Committee on Food, 2003; Thomson, 2004; Rayman, 2004).
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Figures:
Figure 1. The process of brewing Se-enriched beer.
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Figure 2. Production processes of selenized kvass.
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Figure 3. Transformation and bioaccumulation of Se in algae. 1) Selenium uptake via sulfate, phosphate and silicon transporters. 2) Selenium/sulfur assimilation pathway: APSe: adenosine
(seleno)methionine;
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phosphoselenate; GSH: glutathione; OAS: O-acetylserine; (Se)Cys: (seleno)cysteine; (Se)Met: MMT:
dimethyl(di)selenide
methyltransferase; (volatile);
SMT:
DMSeP: selenocysteine
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dimethylselenoproprionate;DM(D)Se:
methylmethionine
methyltransferase. 3) SeCys is incorporated in essential Se-proteins that play a key role in ROS scavenging and defense systems. 4) However, if intracellular Se concentration is high, Se amino acids can also be incorporated into other proteins non-specifically, thus producing malformed structures (Schiavon et al., 2017). Modified with permission from ref 4390660547884 (Elsevier, 2017)
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Figure 4. The production of Se-enriched Tom-Brown flour.
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Figure 5. Comparing toxicity levels of the various form of selenium (sodium selenite and selenium-enriched yeast). Modified with permission from ref 4392380281814 (Cambridge University Press, 2006).
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Graphical abstract
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Organic selenium (Se) is required to sustain proper health in both animals and humans due to its linkage with various biological functions in the immune system.
Some regions around the world are naturally deficient in soil Se whilst others are becoming toxic. Organic Se is more bioavailable and less toxic than the inorganic form.
Studies have shown that the consumption of Se enriched food and supplements leads to
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Supplementing wort with inorganic Se resulted in brewing selenized beverages (beer and
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Kvass).
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the decrease of selenosis.
Figure 1
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Figure 3
Figure 4
Figure 5