Biogeochemistry of selenium and its impact on food chain quality and human health

In areas where soils are low in bioavailable selenium (Se), potential Se deficiencies cause health risks for humans. Though higher plants have been considered not to require this element, the experience with low-Se soils in Finland has provided evidence that the supplementation of commercial fertilizers with sodium selenate affects positively not only the nutritive value of the whole food chain from soil to plants, animals and humans but also the quantity of plant yields. The level of Se addition has been optimal, and no abnormally high concentrations in plants or in foods of animal origin have been observed. Se levels in serum and human milk indicate that the average daily intake has been within limits considered to be safe and adequate. In fact, plants act as effective buffers, because their growth is reduced at high Se levels. They also tend to synthesize volatile compounds in order to reduce excess Se. On the other hand, when added at low concentrations, Se exerts a beneficial effect on plant growth via several mechanisms. As in humans and animals, Se strengthens the capacity of plants to counteract oxidative stress caused by oxygen radicals produced by internal metabolic or external factors. At proper levels it also delays some of the effects of senescence and may improve the utilization of short-wavelength light by plants. High additions are toxic and may trigger pro-oxidative reactions. Thus, the present supplementation of fertilizers with Se can be considered a very effective and readily controlled way to increase the average daily Se intake nationwide.     

Effects of nationwide addition of selenium to fertilizers on foods,and animal and human health in Finland: From deficiency to optimal selenium status of the population

Despite different geological features the Nordic countries are generally selenium-poor areas. In each country various factors such as food importation and life-style determine the selenium (Se) intake. Due to an extremely low Se intake in the 1970s in Finland, 0.025 mg/day, an official decision was made in 1984 to supplement multinutrient fertilizers with Se in the chemical form of sodium selenate. Almost all fertilizers used in Finland since 1985 have contained Se. Currently all crop fertilizers contain 15 mg Se/kg. Finland is still the only country to take this country-wide measure.In a national monitoring programme, sampling of cereals, basic foodstuffs, feeds, fertilizers, soils, and human tissues has been carried out annually since 1985 by four governmental research organizations. Sampling of foods has been done four times per year and human blood has been obtained annually from the same (n = 60) adults. The accuracy of analyses has been verified by annual interlaboratory quality control. During this programme the selenium concentration of spring cereals has increased on average 15-fold compared with the level before the Se fertilization. The mean increase in the Se concentration in beef, pork and milk was 6-, 2- and 3-fold. In terms of Se, organically grown foods of plant origin are generally comparable to products produced before the Se supplementation of fertilizers. Milk from organically fed cows is 50% lower in Se than the usual milk. The average dietary human intake increased from 0.04 mg Se/day/10 MJ in 1985 to a present plateau of 0.08 mg Se/day/10 MJ, which is well above the current nutrition recommendations. Foods of animal origin contribute over 70% of the total daily Se intake. The mean human plasma Se concentration increased from 0.89 μmol/L to a general level of 1.40 μmol/L that can be considered to be an optimal status. The absence of Se deficiency diseases and a reference population have made conclusions on the impact on human health difficult. However, the rates of cardiovascular diseases and cancers have remained similar during the pre- and post-supplementation indicating medical and life-style factors to be much stronger determinants than Se. The nationwide supplementation of fertilizers with sodium selenate is shown to be effective and safe in increasing the Se intake of the whole population. Also, the health of animals has improved.     

Performance,carcass characteristics and meat quality of Nellore cattle supplemented with supranutritional doses of sodium selenite or selenium-enriched yeast

The enrichment of meat with selenium is important to improve the intake of selenium by humans. The effects of supranutritional doses of sodium selenite or selenium-enriched yeast on performance, carcass characteristics and meat quality were evaluated using 63 Nellore cattle in a completely randomized design with two sources (sodium selenite and selenium-enriched yeast), three levels (0.3, 0.9 and 2.7 mg Se/kg DM) and control treatment (without addition of selenium). Final body weight (BW), average daily gain, dry matter intake and gain to feed ratio (G : F) at the end of 84 days of supplementation were not influenced by treatments (P>0.05). Values of pH, ribeye area, back fat thickness and marbling score were also not influenced by treatments (P>0.05). Dressing percentage was greater (P=0.02) in Nellore cattle supplemented with organic Se (58.70%) compared to animals supplemented with inorganic Se (57.94%). Hot carcass weight increased (P=0.05) with the increasing of Se levels in the diet. Colour, shear force (SF), cooking and drip loss remained unchanged (P>0.05); however thiobarbituric acid reactive substances was 15.51% higher with inorganic Se compared with organic Se. The selenium concentration in the meat of animals receiving organic selenium was higher (P<0.001) than that of animals receiving sodium selenite, at all levels (0.3; 0.9 and 2.7 mg/kg DM). The meat of animals receiving 2.7 mg of organic Se/kg of DM presented concentration of 372.7 μg Se/kg in the L.dorsi muscle, and the intake of 150 g of this meat by humans provides approximately 100% of the recommended Se intake (55 μg Se/day for adults). Therefore, the use of supranutritional doses of 2.7 mg Se/kg of DM, regardless of source, is a way of naturally producing selenium-enriched meat without compromising performance, carcass characteristics and quality of Nellore bovine meat.     

Selenium increases seed production in Brassica

Selenium (Se) is essential for humans and animals but is not considered to be essential for higher plants. Although researchers have found increases in vegetative growth due to fertiliser Se, there has been no definitive evidence to date of increased reproductive capacity, in terms of seed production and seed viability. The aim of this study was to evaluate seed production and growth responses to a low dose of Se (as sodium selenite, added to solution culture) compared to very low-Se controls in fast-cycling Brassica rapa L. Although there was no change in total biomass, Se treatment was associated with a 43% increase in seed production. The Se-treated Brassica plants had higher total respiratory activity in leaves and flowers, which may have contributed to higher seed production. This study provides additional evidence for a beneficial role for Se in higher plants.     

Lack of an effect of dairy protein (casein) and soy protein on plasma cholesterol of strict vegetarians. An experiment and a critical review

In animals, ingestion of casein, the principal protein in milk, causes hypercholesterolemia, whereas in humans this effect has not been documented. We added 27 g of casein (the amount in 1.1 liters of skim milk and nearly twice the average U.S. intake) for 20 days, and 27 g of soy protein for an additional 20 days to the daily diet of 13 strict vegetarians who consumed no other animal protein during the study period. The protein supplementation increased the ad libitum daily protein intake from 59 g to 82 g. Levels of plasma LDL, HDL, and total cholesterol were not significantly affected by either the casein or the soy supplementation. Over the 40 days of protein supplementation, there were progressive decreases in VLDL cholesterol (VLDL-C) and increases in triglycerides (TG) from pre-study levels, demonstrated by an overall change in the VLDL-C/TG ratio from 0.30 to 0.17 (P = 0.003). Caloric intake and body weight did not change significantly. From the literature on dietary protein and blood lipid levels and from the present data, it appears that neither the amount of protein in the diet nor whether the protein comes from animal or vegetable sources has an important effect on plasma LDL and HDL levels in humans when consumed in physiologic amounts.     

Bioavailability and possible benefits of wheat intake naturally enriched with selenium and its products

Bioavailability and possible benefits of wheat intake naturally enriched with selenium and its products was tested. Wheat obtained by application of an original combination and procedure for foliar supplementation of plants with Se was characterized on the average by five times higher content of Se, the main form being l-(+)-selenomethionine (SeMet). Substitution of Se-deficient wheat by wheat naturally enriched with Se and its products contributed to the increase of daily intake on the average by 18 μg (12–35 μg) in volunteers, which is more than 50% of the average daily intake. Six weeks after the beginning of its application, increased daily intake of Se brought about the increase of its concentration in the plasma of the examined persons by 53%, in their erythrocytes by 37%, in their hair by 44%, and in their urine by 54%. This result was comparable to the effect obtained in the course of an 8-wk daily intake of supplements with 100 μg Se in the form of enriched bakery yeast. Analysis of glutathione peroxidase (GSH-Px) activity in blood, thiobarbituric acid reactive substances (TBARS) in plasma, lipid parameters (total cholesterol, high-density lipoprotein cholesterol, low-density lipoprotein cholesterol, and triglycerides), and glucose in serum of volunteers showed that the increased Se intake induced increased GSH-Px activity in blood and decreased concentrations of TBARS, lipid parameters, and glucose in blood. Using only one crop (wheat enriched with Se), the existing deficiency of Se in our population can be alleviated. In this way, one-fourth of our population with lower Se intake than 21 μg/d will satisfy basal requirements, whereas one-half will become moderately deficient in Se instead of distinctly deficient in Se.     

The green technology of selenium phytoremediation

Selenium toxicity is encountered in arid and semi-arid regions of the world with alkaline, seleniferous soils derived from marine sediments. Once present in soils and waters at high concentrations, Se is very complicated and highly expensive to remove with conventional physical and chemical techniques. Phytoremediation is a plant-based technology that is being considered for managing Se in central California soils. The technology involves the use of plants in conjunction with microbial activity associated with the plants to extract, accumulate, and volatilize Se. Once absorbed by plant roots, Se is translocated to the shoot where it may be harvested and removed from the site. Therefore, plant species used for phytoremediation of Se-laden soils may by plant uptake and volatilization minimize the Se load eventually entering agricultural effluent and the harvested crop can be carefully blended with animal forage and fed to animals in Se-deficient areas.     

Evaluation of the Potential of Peas (Pisum sativum L.) to Be Used in Selenium Biofortification Programs Under Mediterranean Conditions

Selenium (Se), which has antioxidant, anticancer, and antiviral properties, is an essential micronutrient for humans and animals. This micronutrient is found in high quantity in legumes. Peas have an ever-increasing importance in Spain, and to increase their nutritional value, two foliar Se fertilizers: sodium selenate and sodium selenite, at five different rates: 0, 10, 20, 40, 80 g?ha^?1, were studied during the 2010/2011 crop season on semiarid Mediterranean conditions. Sodium selenate was much more effectively taken up by plants compared to sodium selenite. There was a strong linear relationship between the total Se content and Se rate in both sodium selenate and selenite. For each gram of Se fertilization as either sodium selenate or sodium selenite, the increase of total Se concentration in the grain was 148 and 19 μg Se?kg^?1 dry weight, respectively. Ingestion of 100 g of peas previously fertilized with 10 g of sodium selenate per hectare would result in an intake of 179 μg of Se. This is almost 90 % of the daily recommended dose needed to reduce the chance of some cancers and about 179 % of the minimum concentration required to prevent Se deficiency diseases in animals. The pea has shown to have a strong ability to uptake and accumulate Se under Mediterranean conditions; therefore, this would make it a very strong candidate for inclusion in biofortification programs aiming to increase Se in the food chain.     

The influence of dietary selenium and vitamin E on glutathione peroxidase and glutathione in the rat

The effect of dietary selenium (Se) and vitamin E supplementation on tissue reduced glutathione (GSH) and glutathione peroxidase activity has been studied in the rat. Increasing Se intake by 0.4 ppm gave significantly higher enzyme levels in all tissues studied, an effect not influenced by vitamin E intake. Further increasing Se to 4 ppm gave higher enzyme levels in red blood cells only, while in liver was there was a significant decrease in enzyme activity probably reflecting Se hepatotoxicity. In the absence of Se supplements increasing dietary vitamin E to 100 mg/kg diet significantly increased enzyme activity but this effect was modified by simultaneous Se supplementation.Se intake had no effect on GSH levels. Rats on high vitamin E intake 500 mg/kg had a significantly higher tissue GSH level. Dietary Se had a sparing effect on vitamin E, rats supplemented with Se having significantly raised plasma vitamin E levels.These results confirm the role of selenium in glutathione peroxidase and also show that vitamin E influences the activity of the enzyme.     

Interactions between selenium and iodine uptake by spinach (Spinacia oleracea L.) in solution culture

Interactions between Se (as selenate) and I (as iodate) uptake by spinach plants (Spinacia oleracea L.) were studied under controlled conditions using solution culture. Spinach readily accumulated both Se and I in the edible parts, the leaves, with solution-to-leaf transfer factors ranging from 3.5 to 13.4. The distribution coefficients between leaves and roots ranged from 4.07 to 5.66 for I and 4.51 to 8.59 for Se. Selenium concentrations in plant tissues were unaffected by addition of I to the nutrient solution. Similarly, plant I concentrations were unaffected by addition of Se to the nutrient solution, except in nutrient solution with I at a concentration of 50 μM, in which addition of Se lowered shoot I concentrations significantly, but the effect was of low magnitude. These results indicate the possible feasibility of dual supplementation of plant growth substrates with Se and I to improve human nutrition where these two elements are deficient in the diet. The data also indicate the involvement of a positive feedback mechanism in the uptake of Se by spinach plants, since Se concentrations in leaves increased disproportionately with increasing Se concentration in the nutrient solution.     

Microbial-enhanced Selenium and Iron Biofortification of Wheat (Triticum aestivum L.) - Applications in Phytoremediation and Biofortification

Selenium (Se) is an essential trace element for humans and other mammals. Most dietary Se is derived from crops. To develop a Se biofortification strategy for wheat, the effect of selenate fertilization and bacterial inoculation on Se uptake and plant growth was investigated. YAM2, a bacterium with 99% similarity to Bacillus pichinotyi, showed many plant growth promoting characteristics. Inoculation with YAM2 enhanced wheat growth, both in the presence and absence of selenate: YAM2-inoculated plants showed significantly higher dry weight, shoot length and spike length compared to un-inoculated plants. Selenate also stimulated wheat growth; Un-inoculated Se-treated plants showed a significantly higher dry weight and shoot length compared to control plants without Se. Bacterial inoculation significantly enhanced Se concentration in wheat kernels (167%) and stems (252%), as well as iron (Fe) levels in kernels (70%) and stems (147%), compared to un-inoculated plants. Inoculated Se-treated plants showed a significant increase in acid phosphatase activity, which may have contributed to the enhanced growth. In conclusion; Inoculation with Bacillus sp. YAM2 is a promising Se biofortification strategy for wheat and potentially other crops.     

Plant-Associated Symbiotic Burkholderia Species Lack Hallmark Strategies Required in Mammalian Pathogenesis

Burkholderia is a diverse and dynamic genus, containing pathogenic species as well as species that form complex interactions with plants. Pathogenic strains, such as B. pseudomallei and B. mallei, can cause serious disease in mammals, while other Burkholderia strains are opportunistic pathogens, infecting humans or animals with a compromised immune system. Although some of the opportunistic Burkholderia pathogens are known to promote plant growth and even fix nitrogen, the risk of infection to infants, the elderly, and people who are immunocompromised has not only resulted in a restriction on their use, but has also limited the application of non-pathogenic, symbiotic species, several of which nodulate legume roots or have positive effects on plant growth. However, recent phylogenetic analyses have demonstrated that Burkholderia species separate into distinct lineages, suggesting the possibility for safe use of certain symbiotic species in agricultural contexts. A number of environmental strains that promote plant growth or degrade xenobiotics are also included in the symbiotic lineage. Many of these species have the potential to enhance agriculture in areas where fertilizers are not readily available and may serve in the future as inocula for crops growing in soils impacted by climate change. Here we address the pathogenic potential of several of the symbiotic Burkholderia strains using bioinformatics and functional tests. A series of infection experiments using Caenorhabditis elegans and HeLa cells, as well as genomic characterization of pathogenic loci, show that the risk of opportunistic infection by symbiotic strains such as B. tuberum is extremely low.     

Nephrotoxicity and neurotoxicity in humans from organogermanium compounds and germanium dioxide

There is no known biological requirement for germanium (Ge), germanates, or any organogermanium compound. Ge deficiency has not been demonstrated in any animal. The estimated average dietary intake of Ge in humans is 1.5 mg/d. Ge is widely distributed in edible foods, all of which, with few exceptions, contain less than 5 ppm Ge, since higher levels are toxic to most plants. Ingestion of Ge compounds has been shown to produce toxic effects in experimental animals. In recent years inorganic germanium salts and novel organogermanium compounds, such as carboxyethyl germanium sesquioxide (Ge-132) and lactate-citrate-germanate (Ge lactate citrate) have been sold as "nutritional supplements" in some countries for their purported immunomodulatory effects or as health-producing elixirs, resulting in intakes of Ge significantly exceeding the estimated average dietary intake. Since 1982, there have been 18 reported cases of acute renal dysfunction or failure, including two deaths, linked to oral intake of Ge elixirs containing germanium dioxide (GeO2) or Ge-132. In these cases, biopsies show vacuolar degeneration in renal tubular epithelial cells, without proteinuria or hematuria, in the absence of glomerular changes. Serum creatinine levels have been well above 400 mumol/L in such patients. In 17 of 18 cases, accumulated elemental Ge intakes reportedly ranged between 16 to 328 g over a 4-36 mo period, or between 100 to 2000 times the average estimated dietary intake for human. In surviving patients, renal function improved after discontinuation of Ge supplementation. However, in no case was recovery complete. One organogermanium compound, an azaspiran organogermanium compound, 2-aza-8-germanspiro[4,5] decane-2-propamine-8,8-diethyl-N,N-dimethyl dichloride (spirogermanium), has been found to cause both neurotoxicity and pulmonary toxicity in phase I and II studies examining its chemotherapeutic potential as an antitumor drug in the treatment of various malignancies. In cancer patients given the drug spirogermanium, 40% experienced marked, yet transient neurotoxicity. Two patients suffered from pulmonary toxicity. Results of phases I and II human cancer trials for spirogermanium have not been favorable, with the exception of moderate benefits for three types of malignancies. It is recommended that patients exposed to long-term (greater than 3 mo) Ge supplementation at levels well above the estimated daily intake be medically supervised and monitored for potential renal-, pulmonary- or neurotoxicity. Further study regarding the mechanism of Ge-induced nephrotoxicity in human is warranted.     

Selenium and asthma

Se is a potent nutritional antioxidant important for various aspects of human health. Because asthma has been demonstrated to involve increased oxidative stress, levels of Se intake have been hypothesized to play an important role in the pathogenesis of asthma. However, significant associations between Se status and prevalence or severity of asthma have not been consistently demonstrated in human studies. This highlights both the complex etiology of human asthma and the inherent problems with correlative nutritional studies. In this review, the different findings in human studies are discussed along with results from limited intervention studies. Mouse models of asthma have provided more definitive results suggesting that the benefits of Se supplementation may depend on an individual’s initial Se status. This likely involves T helper cell differentiation and the mechanistic studies that have provided important insight into the effects of Se levels on immune cell function are summarized. Importantly, the benefits and adverse effects of Se supplementation must both be considered in using this nutritional supplement for treating asthma. With this in mind new approaches are discussed that may provide more safe and effective means for using Se supplementation for asthma or other disorders involving inflammation or immunity.     

Antioxidative and growth-promoting effect of selenium on senescing lettuce

In human and animal cells, Se plays an essential role in antioxidation and exerts an antiaging function but it is toxic at high dietary intake. To increase its intake in forage and foodstuffs, Se fertilization is adopted in some countries where soils are low in bioavailable Se, even though higher plants are regarded not to require Se. To test its ability to counteract senescence-related oxidative stress in higher plants, a pot experiment was carried out with lettuce (Lactuca sativa) cultivated with increasing amounts of H₂SeO₄. The yields harvested 7 or 14 weeks after sowing revealed that a low Se dosage (0.1 mg kg^–1 soil) stimulated the growth of senescing seedlings (dry weight yield by 14%) despite a decreased chlorophyll concentration. The growth-promoting function was related to diminished lipid peroxidation. In young and senescing plants, the antioxidative effect of Se was associated with the increased activity of glutathione peroxidase (GSH-Px). In the senescing plants, the added Se strengthened the antioxidative capacity also by preventing the reduction of tocopherol concentration and by enhancing superoxide dismutase (SOD) activity. When no Se was added, tocopherols and SOD activity diminished during plant senescence. The higher Se dosage (1.0 mg kg^–1 soil) was toxic and reduced the yield of young plants. In the senescing plants, it diminished the dry weight yield but not the fresh weight yield.     

Effect of selenium on distribution of macro- and micro-elements to different tissues during wheat ontogeny

Selenium (Se) is essential for health of humans, animals, and plants. Especially wheat is a major source of Se in the terrestrial food chain. In this study, an element analysis was optimized and the content of Ca, Mg, K, S, P, Fe, Se, Mn, Cu, Zn, and Mo in leaves, roots, and seeds were measured during growth of wheat (Triticum aestivum L. cv. Manu) in Hoagland nutrient solution with 5 and 15 μM Na₂SeO₄. Se was transported to all investigated tissues and accumulated in the seeds in proportion to used amounts. The supplementation of Se, independently of concentration, weakly modified the micro- and macro-elements content in the seedlings. In the flag-leaf stage, an increase of the Mo and S content in the shoots and the S and Cu content in the roots was found. Moreover, in the generative phase, a decrease in Ca and Fe in the roots was registered. Increased Se in the nutrient solution strongly stimulated the Se accumulation in the seeds.     

Silicon: A neglected micronutrient essential for bone health

Bone matrix is predominantly made up of collagen, and in vitro and in animal models studies have shown that silicon is linked to glycosaminoglycans and plays an important role in the formation of cross-links between collagen and proteoglycans, determining the beneficial effects on strength, composition, and mechanical properties of bone. However, there are still no precise indications regarding a possible role of silicon on bone health in humans. Given this background, the aim of this narrative review was to consider the effectiveness of silicon dietary intake and silicon dietary supplementation (alone or with other micronutrients), in order to suggest a daily dosage of Si supplementation, on bone mineral density in humans. This review included eight eligible studies: four regarding dietary intake and four considering supplementation with silicon alone or with other nutrients. Despite the number of studies considered being low, the number of subjects studied is high (10012) and the results are interesting. Although to date the available scientific evidences are not considered valid enough to allow to establish an adequate level of Silicon intake, based on extrapolations from the data obtained with studies on animal and human models, it has been suggested that an adequate intake in order to promote beneficial effects for bone could be considered to be around 25 mg silicon/day. As for silicon dietary supplements, it has been shown that the combined treatment with orthosilicic acid (6 mg), calcium, and vitamin D has a potentially beneficial effect on femoral BMD compared to only use of calcium and vitamin D.     

Relevance, essentiality and toxicity of trace elements in human health

The metals Mn, Fe, Cu, and Zn, and the non-metal Se are considered "trace elements" (TE) because of their essentiality and very limited quantity in humans. The biological activities of Cu, Fe, Mn, and Se are strongly associated with the presence of unpaired electrons that allow their participation in redox reactions. In biological systems these metals are mostly bound to proteins, forming metalloproteins. Many of the metals in metalloproteins are part of enzymatic systems, have structural and storage functions, or use the protein to be transported to their target site in the organism. In humans Mn, Fe, Cu, Zn, and Se accomplish decisive functions to maintain human health. Deficiency in any of these TE leads to undesirable pathological conditions that can be prevented or reversed by adequate supplementation. In sufficiently nourished persons, supplementation should be carefully controlled, given the toxic effects ascribed to TE when present in quantities exceeding those required for accomplishing their biological functions. The dietary reference intakes provided by national regulatory agencies are guides to define intake, supplementation and toxicity of Mn, Fe, Cu, Zn, and Se, as well other elements considered micronutrients for humans.