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Biogeochemical behavior of selenium in soil-air-water environment and its

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Article in International journal of Environmental Science and Technology · September 2023

DOI: 10.1007/s13762-023-05169-0

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International Journal of Environmental Science and Technology
https://doi.org/10.1007/s13762-023-05169-0

REVIEW

Biogeochemical behavior of selenium in soil‑air‑water environment

and its effects on human health
Z. N. Xu1 · Z. Q. Lin2 · G. S. Zhao1 · Y. B. Guo1

Received: 4 August 2022 / Revised: 3 June 2023 / Accepted: 16 August 2023

© The Author(s) under exclusive licence to Iranian Society of Environmentalists (IRSEN) and Science and Research Branch, Islamic Azad University 2023

Abstract
Selenium as an essential trace element for human beings and animals plays a critical role in human and animal health via
various selenoproteins. Recently, the biogeochemical behaviour of selenium and its relation on human health have received
increasing attention. This review provides a comprehensive understanding of selenium biogeochemical cycling in eco-
systems. The concentration, transport, and transformation of selenium in environments and their effects on human health
are reviewed. Several important aspects of selenium in the environment are detailed mentioned, including (1) effects of
selenium on human health, (2) biogeochemical cycling of selenium in lithosphere, pedosphere, hydrosphere, atmosphere,
and biosphere, (3) metabolic and transport of selenium in organisms. It is emphasized that the biogeochemical behaviors
of selenium in rocks, soil, water, atmosphere, and organisms will impact human selenium intake through the food chain. In
addition, future scientific issues and research attention have also been explored from the selenium biochemical perspectives.

Keywords Selenium · Transport and fate · Dietary intake · Human health · Biogeochemical processes

Introduction through Se-containing amino acid, selenocysteine (SeCys)

Selenium (Se) is a metalloid trace element first detected
in 1817 by Jons Jacob Berzelius. Selenium has been con-
sidered harmful to human health until 1957 when it was
demonstrated to alleviate liver necrosis (Schwarz and Foltz
1957). Selenium is a vital element of glutathione peroxidase
in performing antioxidant functions (Rotruck et al. 1973).
In China, Keshan disease has been effectively mitigated by
Se supplementation (Zhou et al. 2018; Zhu et al. 2019). In
1973, based on the results of Se in human health, the World
Health Organization (WHO) announced Se as an indispen-
sable nutrient for humans. From then, the function of Se on
human various physiological activities were identified. Sele-
nium is important for some biological functions in humans
Editorial responsibility: Samareh Mirkia.
* Y. B. Guo
guoyb@cau.edu.cn
1
College of Resources and Environmental Sciences, China
Agricultural University, Beijing 100193, China
2
Department of Environmental Sciences & Department
of Biological Sciences, Southern Illinois University,
Edwardsville, IL 62026‑1099, USA
that forms different selenoproteins (Avery and Hoffmann
2018; Guillin et al. 2019). In the human body, approximately
80% of Se is used for the synthesis of selenoproteins, which
were essential for many biological processes (Minich 2022).
Therefore, Se deficiency has been related to the etiology of
human diseases (Dinh et al. 2018; Rayman 2020). In human
physiology, Se is ingested mainly from food sources, and
the Se content in dietary materials will directly or indi-
rectly be determined by the Se concentration and speciation
in agricultural soil. The uneven soil Se distribution might
contribute to differences in human Se dietary intake around
the world. Crops and livestock in some countries have been
reported to have Se deficiencies to varying degrees, affect-
ing about 15% of the world's population (Dinh et al. 2018;
Fordyce 2013; White et al. 2012).
Selenium has been widely used in various fields such
as electronics, metallurgy, agriculture, and medicine (U.S.
Geological Survey 2021). In recent years, Se contamina-
tion and remediation techniques (He et al. 2018a; Okonji
et al. 2021; Schiavon and Pilon-Smits 2017), microbial
Se reduction and resistance (Nancharaiah and Lens 2015;
Nie et al. 2023; Tugarova and Kamnev 2017; Wang et al.
2022a), Se intake and human health (Ekumah et al. 2021;
Rayman 2020), mechanisms of Se uptake, metabolism and

13
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hyperaccumulation in plants (Lima et al. 2018; White 2016,
2018), Se cycling in soil-microbe-plant systems (Guo et al.
2023; Kushwaha et al. 2021; Qu et al. 2023; Uallh et al.
2019; Wang et al. 2022b; Winkel et al. 2015) and particu-
larly Se biofortification of food crops (Hossain et al. 2021;
Sarwar et al. 2020) have been reviewed by multiple research-
ers. However, the effects of environmental distributions of
Se and Se biogeochemical processes on human health have
not been addressed comprehensively. The biogeochemical
cycling of Se plays an important role in the distribution of
Se in the environment, which ultimately affects the intake
of Se in humans (Blazina et al. 2014). Thus, this review
focused on Se transport pathways, transformation, fate, and
distribution among different environmental components in
relation to their impacts on Se biofortification, human Se
dietary intake, and ultimately on human health.
Selenium as essential nutrient for human
health
Selenium dietary intake
For humans, Se comes from the daily intake of water, ani-
mal, and plant foods, and the content of Se in water and
foods is variety in different regions of the world. Plant-
derived foods provide the principal source of Se for most
human populations. Rice, wheat, and corn are the three pil-
lars of the human diet and are closely related to human nutri-
tion (Neumann et al. 2010). Over half of the world's popu-
lation consumes rice as a major staple food (HarvestPlus
2022). The Se content in rice is 0.022 μg ­g−1 in Argentina,
0.10 μg ­g−1 in Brazil, 0.019 μg ­g−1 in Greece, 2.01 μg ­g−1 in
Nigeria, and in China, it varies from 0.0021 to 2.11 μg ­g−1
(Dinh et al. 2018; Gbadebo et al. 2010; Lemire et al. 2010;
Pappa et al. 2006; Sigrist et al. 2012). For the majority of
developing countries, wheat contributes approximately 50%
of the daily caloric intake, and for rural areas, wheat con-
tributes more than 70% of caloric intake (Cakmak 2008).
The Se content of wheat in China varies from 0.0052 to
0.44 μg ­g−1, and in Algeria, the Se content of wheat dif-
fers from 0.021 to 0.153 μg ­g−1 (Beladel et al. 2013; Dinh
et al. 2018). Meanwhile, animal-based foods and seafood
are the best providers of Se for humans, and their Se content
exceeds that of plant food (Dinh et al. 2018). In Greece,
the Se levels of pork (0.094 μg ­g−1) was higher than that
of chicken (0.079 μg ­g−1) and beef (0.049 μg ­g−1) (Pappa
et al. 2006). In Croatia, Se content ranges from 0.036 to
0.098 μg ­g−1 in pork and 0.035 to 0.15 μg ­g−1 in beef. The
Se content in kidney is the highest, which is 1.72 μg ­g−1 in
beef kidney and 1.67 μg ­g−1 in pork kidney, respectively
(Bilandzic et al. 2021). In Pakistan, the Se content of mutton
is 0.11–0.13 μg ­g−1, beef is 0.083–0.099 μg ­g−1, and chicken
13
International Journal of Environmental Science and Technology
is 0.082–0.098 μg ­g−1 (Almani et al. 2020). As for seafood,
take Atlantic salmon as an example, which is one of the
most popular seafood, with Se content ranging from 0.009 to
0.034 μg ­g−1 (Moxness et al. 2022). Oysters have the highest
Se content in the "shellfish" food group, with the content
ranging from 1.66 to 4.88 μg ­g−1 (Bilandzic et al. 2014).
There are about 20 mg Se accumulated in the human body
of 70 kg, with 27.5–46.9% in skeletal muscle, 15.9% in bone,
7.6–10.2% in blood, and 2.8–3.6% in kidney (Combs 2013;
Oster et al. 1988; Zachara et al. 2001). In healthy humans,
Se concentrations are highest in the kidneys, followed by the
liver, spleen, pancreas, heart, brain, lung, bone, and skeletal
muscle (Oster et al. 1988; Zachara et al. 2001). The main
source of Se in the human body is dietary intake, primarily
from plant-based foods (El-Ramady et al. 2015; Institute
of Medicine, Food and Nutrition Board 2021). Human Se
intake is related to dietary structure and Se content in food,
which leads to the variety of Se intake in countries and
regions. The daily Se intake of Chinese people ranges from
10.96 to 2144 μg (Dinh et al. 2018). The daily Se intake of
Mumbai adults is 61.9 μg, while that of Italy is 50.9 μg,
and that of pregnant women (1003 subjects, 20–40 years
old) in the United States is 110–114 μg (Amodio-Cocchieri
et al. 1995; Bailey et al. 2019; Mahapatra et al. 2001). In
Croatia, the daily intake of Se is 37.8 μg for local women
(61 subjects, 20–60 years old), 20–53 μg for people liv-
ing in Turkey (40 subjects), 17.1 μg for children (159 sub-
jects, 6–59 months) and 36.1 μg for women (111 subjects,
19–39 years old) in central highlands of Kenya (Aras et al.
2001; Matek et al. 2000; Ngigi et al. 2020). In Irish, the
daily intake of Se in the total population (1500 subjects,
18–90 years old) is 71.7 μg, 104.9 μg for adults in Madaba
(Jordan, 500 subjects, 18–60 years old), 93.2 μg for people
living in Riyadh (Saudi Arabia, 260 subjects), 109 μg for
Newfoundland population (2420 subjects), 51 μg for ado-
lescent girls in Iceland (96 subjects, 16–20 years old), and
84.3–105.9 μg for adolescents in Brazil (76,957 subjects,
12–17 years old) (Al-Othman et al. 2012; Buffini et al. 2023;
Gudmundsdottir et al. 2012; Hammouh et al. 2020; Retond-
ario et al. 2019; Wang et al. 2017a).
As an essential nutrient element, Se can incorporate into
at least 25 proteins through Se-containing amino acid, SeCys
(Avery and Hoffmann 2018; Guillin et al. 2019). Seleno-
proteins are mainly classified as glutathione peroxidases
(GSH-Px), thioredoxin reductases (TrxR), and deiodinases
(Dio) (Ekumah et al. 2021; Pitts et al. 2014). There are many
other selenoproteins in the human body, which are named
with the prefix seleno-, such as selenoprotein K (SelK),
selenoprotein S (SelS), and selenoprotein P (SelP) (Ekumah
et al. 2021). Earlier studies have demonstrated that some
selenoproteins such as GSH-Px, SelK, SelS, SelP, and TrxR
can be involved in antioxidation, cell signaling, enhancing
immunity, and thyroid hormone production (Avery and
International Journal of Environmental Science and Technology
Hoffmann 2018; Chauhan et al. 2019; Guillin et al. 2019).
In human body, Se is transported primarily by SelP, which
is significantly correlated with the activity and number of
pancreatic β-cells (Burk and Hill 2015; Saito 2020). Sele-
noprotein S participates in the endoplasmic reticulum (ER)
membrane-localized multiprotein complex and is involved
in the degradation of misfolded proteins, and it has a high
level of expression in skeletal muscle with the potential
function to regulate contractile skeletal muscle (Addinsall
et al. 2018). In addition, SelK is also involved with the ER
membrane-localized multiprotein complex that regulates cell
­Ca2+ flux and influences T cell proliferation and differentia-
tion (Shchedrina et al. 2011; Wang et al. 2017b). The pres-
ence of excessive amounts of Se in the body could be toxic,
even though Se is necessary for health. There is a relatively
narrow gap between Se deficiency and toxicity. Overall,
human Se poisoning has been only observed with a limited
number of incidents and/or in seleniferous areas, while Se
deficiency could result in adverse health effects and becomes
a major public health issue worldwide (Dinh et al. 2018;
Rayman 2020). The low Se areas are generally considered
as the region with soil Se levels lower than 0.125 mg ­kg−1,
while the region with soil Se content exceeding 3.0 mg ­kg−1
belongs to the Se excess region (Tan 1989). The concentra-
tions of Se in edible plant tissues are often related to the soil
Se content, and thus, human daily dietary intake of Se can
be affected significantly by soil Se contents in the environ-
ment (El-Ramady et al. 2015; Sarwar et al. 2020). For the
sake of human health, daily Se dietary intake needs to be
regulated. For example, 55 μg ­d−1 per capita (adult) was rec-
ommended in the U.S., 60 μg ­d−1 per capita (adult) in China,
60–70 μg ­d−1 per capita (adult) in Germany, Austria, and
Switzerland, and 70 μg ­d−1 per capita (adult) in Australia
and New Zealand, while the tolerable upper intake level was
400 μg ­d−1 per capita (adult) (Kipp et al. 2015; Institute of
Medicine, Food and Nutrition Board 2021; National Health
and Medical Research Council 2006; National Health Com-
mission of the People’s Republic of China 2017).
Effects of low selenium environment and Se intake
on human health
The amount of bioavailable Se in the environment ulti-
mately determines the quantity of Se available for human
intake through the abundance of Se in edible plant tissues or
through food chains (Oksanen and Sandholm 1970; Trippe
and Pilon-Smits 2021). Plants provide approximately 60%
of human daily dietary consumption of Se, followed by meat
and seafood. Drinking water and beverages generally provide
small amounts of Se except in seleniferous areas (Kieliszek
2019; Qin et al. 2013). Selenium deficient soils are mainly
formed by weathering of igneous rocks with low Se content
(Schiavon et al. 2020). The existence of Se deficient areas
has been reported in several countries or regions around the
world, such as Briazil, Canada, China, Denmark, Finland,
India, Nepal, New Zealand, North Korea, Peru, and a wide
part of Europe (Dinh et al. 2018; do Nascimento et al. 2021;
Ekumah et al. 2021; Roman 2016; Kieliszek 2019). In Se
deficient areas, low levels of soil Se resulted in low Se con-
tents in food products, resulting in deficient human daily
dietary Se intake compared with the nonendemic control
population (Chen 2012). It was found that the epidemic
outbreaks of Keshan disease and Kashin-Beck disease were
linked to low soil Se in China (Dinh et al. 2018). Prolonged
deficiency of Se in the body will lead to serious diseases.
The deficiency of Se reduces immunity, damages the nerv-
ous system, and has a negative impact on fetal development
(Steinbrenner and Sies 2013). It has a negative influence
on testis development and sperm production (Oldereid et al.
1998; Xu et al. 2023a). Additionally, Alzheimer’s disease
and depression have been associated with Se deficiency (Saj-
jadi et al. 2022; Zhang and Song 2021). Selenium deficiency
can also lead to muscle inflammation, red blood cell fragil-
ity, myocardial dysfunction, kidney disease, Keshan disease,
Kashin-beck disease, and even with increased risk of cancer
(Fordyce 2013; Rayman 2020).
Effects of excessive selenium exposure on human
health
Seleniferous soils were developed mainly from sedimentary
deposits with much higher Se concentrations than igneous
rocks (Dhillon et al. 2019). Excessive Se exposure and
uptake might also have adverse effects on human health in
seleniferous regions, such as Punjab in India, the northern
U.S., and parts of China, Venezuela and Colombia (Chawla
et al. 2020; Dhillon and Dhillon 2014; Rayman 2008). In
Punjab, because of high levels of Se in soil and water, the Se
contents in mustard and wheat plants were 931 mg ­kg−1 and
390 mg ­kg−1, respectively (Eiche et al. 2015). The consump-
tion of those Se-laden crops will increase Se dietary intake
and accumulation in the human body, leading to harmful
impacts or Se poisoning (Chawla et al. 2020). Selenium
can replace sulphur in amino acids (cysteine and methio-
nine) and thus lead to the production of malformed proteins
(Gupta and Gupta 2017; Kolbert et al. 2019). There are a
variety of symptoms of excessive intake or Se poisoning,
including alopecia, nail damage, dermatitis, hypotension,
tachycardia, muscle contractions dizziness, nausea, vomit-
ing, facial flushing, tremors, muscle aches and nervous sys-
tem disorders (Dinh et al. 2018; Hossain et al. 2021; Rayman
2020; Vinceti et al. 2016, 2019; Yang et al. 1983). Exces-
sive Se intake also reduces sperm density and motility, and
also causes testicular damage (Xu et al. 2023a). Studies have
also reported that excessive intake of Se is associated with
diabetes, hypertension and many types of cancer (Vinceti
13

et al. 2018). Acute Se toxicity can cause severe intestinal
problems, heart injury, renal failure and death (Hadrup and
Ravn-Haren 2020). However, the reports of Se toxicity in
humans are limited because the high concentration of foods
and seleniferous environment were unfrequent on earth.
Distribution of selenium in the environment
Selenium in the lithosphere
As the main natural source of Se, the total Se content in
rocks accounts for nearly 40% of Se in Earth crust (Wang
and Gao 2001). The average concentration of Se in Earth
crust is 0.13 mg ­kg−1, ranging from 0.09 to 0.2 mg ­kg−1
in the upper crust to the lower crust of Earth (Rudnick
and Gao 2014). Yang et al. (1983) reported that contents
of Se were up to 84,123 mg ­kg−1 in Se-rich carbonaceous
siliceous rock at the top of the Permian Maokou Forma-
tion in the township of Yutangba in Enshi city, Hubei of
China (Table 1). In a different seleniferous region located
in Ziyang county, Shaanxi of China, concentrations of Se in
the Ediacaran-early Cambrian Se-rich black shale and the
Cambrian carbonaceous siliceous rock were 303 mg ­kg−1
and 278 mg ­kg−1, respectively (Table 1) (Feng et al. 2012;
Long and Luo 2017). High levels of Se were also observed
in other sedimentary rocks like limestones and shales. Ear-
lier studies showed that Se was the most abundant trace ele-
ment in the Early Cambrian black shale, while low levels of
Se were generally found in volcanic rocks (Fordyce 2013;
Tian et al. 2017). As for western U.S. soil, Se comes primar-
ily from Cretaceous shale deposits in the prehistoric inland
sea, which is also the principal source of Se in other Se-rich
areas (Hladun et al. 2013; Tuttle et al. 2014; Wadgaonkar
et al. 2018). Furthermore, different Se minerals have been
observed in natural rocks, commonly berzelianite (­ Cu2Se),
tiemannite (HgSe), and naumannite ­(Ag2Se) (Fordyce 2013;
Wadgaonkar et al. 2018).
Selenium in the pedosphere
Concentrations of Se in natural soil is closely related to the
constitution of its geological parent material, while the soil
Se distribution reflects the soil formation process and partial
levels of atmospheric Se deposition (Table 2) (El-Ramady
et al. 2014). Selenium in soil generally comes from the
process of rock weathering and exists in soil as inorganic
and organic forms. Inorganic Se mainly includes selenate
­(SeO42−), selenite (­ SeO32−), and elemental Se (­ Se0), while
common organic Se compounds are SeCys and selenom-
ethionine (SeMet) (Fordyce 2013). In soil, pH and redox
potential (Eh) contribute significantly to Se chemical forms
(Dinh et al. 2019). The amount of Se in soils varies globally,
13
International Journal of Environmental Science and Technology
most parts of the world soil commonly vary between
0.01 and 2.00 mg ­kg−1, with an average concentration of
0.4 mg ­kg−1, and in seleniferous areas, the soil Se concentra-
tion is up to 86.59 mg ­kg−1 (Enshi, China) and even up to
1200 mg ­kg−1 (Meath, Ireland) (Table 2) (Dinh et al. 2018;
Fleming and Walsh 1957; Li et al. 2020). Concentrations
of Se in soil types of forest soils, organic soil, and calcare-
ous soils are generally high (El-Ramady et al. 2014; Floor
and Román-Ross 2012; Fordyce 2013). According Se con-
centration, soils can be categorized into: Se deficient soil,
Se marginal soil (0.125–0.175 mg ­kg−1), Se moderate soil
(0.175–0.40 mg ­kg−1), Se high soil (0.40–3.0 mg ­kg−1), and
Se excessive soil (Tan 1989).
Selenium in the hydrosphere
In aquatic environments, Se is mainly present in low levels of
selenate and selenite (Table 3) (Winkel et al. 2015). In gen-
eral, Se contents in fresh water are usually below 1 μg ­L−1,
range from 0.02 to 0.5 μg ­L−1, 0.1 to 0.35 μg ­L−1, and 0.06
to 400 μg ­L−1 in river water, sea water, and groundwater,
respectively (Table 3) (El-Ramady et al. 2014). In ground-
water, the primary sources of Se come from rock weather-
ing, soil or rock leaching, excessive rainfall, irrigation in
Se-rich soils, and dissolution of soluble salts (Fig. 1) (Kumar
and Riyazuddinb 2011; Leblanc et al. 2018). The dominant
chemical Se species in water can be significantly affected by
temperature, pH, Eh, microbial activity, and organic matter
content (Kumar and Riyazuddinb 2011; Martin et al. 2018).
Selenium in the atmosphere
The atmosphere is a significant transitory reservoir for
global Se biogeochemical cycling, as well as a significant
contributor to regional or global Se distribution. Biogenic
volatilization of Se from land and ocean surfaces to the
atmosphere involves microbial methylation of Se in the
soil, plant, and water environment. Globally, the emission
of volatile Se from the terrestrial and ocean surfaces is
1.2 × ­106 kg and 5–8 × ­106 kg per year, respectively, while
the emission from agricultural land is 0.1–10 Se ­m−2 ­year−1
(Lin 2019). The global average Se amount in the atmos-
phere is roughly 0.2 ng ­m−3, and the amount of Se in the
atmosphere varies greatly in the world (Table 4) (El-Ramady
et al. 2014). Selenium in the atmosphere could present in
both particulate and gaseous forms, including airborne inor-
ganic Se (e.g., H­ 2Se, ­Se0, and S
­ eO2) and organic gaseous Se
(Fig. 1) [e.g., dimethylselenide (DMSe) and dimentyldisele-
nide (DMDSe)]. Gaseous Se compounds in the atmosphere
can be oxidized by airborne free radicals into Se-containing
aerosols.
International Journal of Environmental Science and Technology

Table 1  Selenium contents in rocks

Country/Region Type of rocks Se contents Location References

Brazil Volcanic rock 0.013–0.24 mg ­Kg−1 Parana Basin Hughes et al. (1986)
China Argillaceous dolomite 0.13–9.94 mg ­Kg−1 Yangtze Gorges area Tian and Luo (2017)
Argillaceous limestone 0.14–0.99 mg ­Kg−1
Black shale 0.35–25.08 mg ­Kg−1
Calcareous shale 0.18–2.45 mg ­Kg−1
Calcilutite 0.09–0.78 mg ­Kg−1
Clay 1.83–30.08 mg ­Kg−1
Dolostone 0.02–1.12 mg ­Kg−1
Limestone 0.04–2.05 mg ­Kg−1
Muddy dolostone 0.02–4.00 mg ­Kg−1
Shale 0.178.05 mg ­Kg−1
Silicalite 0.03–5.45 mg ­Kg−1
Siltstone 0.08–1.23 mg ­Kg−1
Silty shale 0.08–0.12 mg ­Kg−1
Tillite 0.03–0.39 mg ­Kg−1
Black shales (Ediacaran) 0.25−30.09 mg ­Kg−1 Linwu Tian et al. (2017)
Black shales (Early Cambrian) 0.54–5.01 mg ­Kg−1 Gushui
Coal 13–1332 mg ­Kg−1, Enshi Yang et al. (1983)
up 84,123 mg ­Kg−1
Dolomite 0.05–0.07 mg ­Kg−1 Enshi Li et al. (2020)
Bioclastic limestone 0.05–0.15 mg ­Kg−1
Sandstone 0.08–0.10 mg ­Kg−1
Siliceous rock 12.9–50.9 mg ­Kg−1 Huangboshuwan Shuangan Feng et al. (2012)
Carbonaceous siliceous rock 260–278 mg ­Kg−1 Shuangan
Black shale 303 mg ­Kg−1 Southern of South Qinling Moun- Long and Luo (2017)
tains
Dolomistic limestone 85 mg ­Kg−1
Siliceous rock 104 mg ­Kg−1
France Peridotite ­ g−1 Eastern Pyrenees
< 3.5–100.7 μg K Lorand and Alard (2010)
India Claystone, Salt and pepper sand- 1864–2754 μg ­Kg−1 Dhillon and Dhillon (2014)
stone, yellowish brown sandstone,
light grey sandstone
Sandstone, shale, clay-stone, con- 11–847 μg ­Kg−1
glomerates, siltstone, limestone
Sandstones, claystone, siltstones, 46–644 μg ­Kg−1
wood coal
Ireland Shale 14.3–42 mg ­Kg−1 Ballybunion Bay Armstrong et al. (2019)
Shale 9.2–44.4 mg ­Kg−1 Nun's Beach
Shale 9.4–12.2 mg ­Kg−1 Inishcorker
Shale 62.6–93.5 mg ­Kg−1 Whiddy Island
Korea Black shale and slate 1.6–2.3 mg ­Kg−1 Dukpyung Park et al. (2010)
Phyllite and limestone 0.9 mg ­Kg−1
Black shale and slate 1.5–33.9 mg ­Kg−1 Chubu
Nigeria Shale 1.6–5.0 mg ­Kg−1 Lower Benue trough and the Niger Nganje et al. (2020)
Delta Basin
Saudi Arabia Gypsum, shale and sand 0.6–1.5 mg ­Kg−1 Coastal areas of the Gulf of Aqaba, Ghrefat et al. (2016)
Saudi Arab
Conglomerate and sandstone 0.9–1.6 mg ­Kg−1
Hemantic sandstone 1.6 mg ­Kg−1
Conglomerate 0.9 mg ­Kg−1
Sandstone 0.9 mg ­Kg−1

13

Table 1  (continued)
Country/Region Type of rocks Se contents
−1
Granite 1–2 mg ­Kg
Rhyolite porphyritic 0.6 mg ­Kg−1
Metatuff 0.9 mg ­Kg−1
Metavolcanic 0.9 mg ­Kg−1
England Black shale 28.8–116.0 mg ­Kg−1 Porth Felen
Red mudstone < 0.2–0.6 mg K­ g−1 Carreg
Pyritic basalt 0.7–1.2 mg ­Kg−1
Massive sulphide 1.4–1.5 mg ­Kg−1
Coal 0.4–61.9 mg ­Kg−1
Selenium in the biosphere
Selenium in the environment can be absorbed and trans-
formed by microorganisms, plants, and animals. Inorganic
Se can be absorbed by plants and microbes and converted
into organic Se compounds (Harvey et al. 2020; Hu et al.
2019a, b, 2020, 2021; Martinez et al. 2020). Moreover,
organic Se in the environment can also be absorbed by plants
and microbes, and organic Se is absorbed more efficiently by
plants than inorganic Se (Kikkert and Berkelaar 2013; Wang
et al. 2020a; Zhu et al. 2018). Herbivores acquire Se by feed-
ing on plants and then Se is transported in the food chain.
Schuler et al. (1990) observed that Se was biomagnified by
nearly 5,000 times through trophic levels at a marshland in
Central California.
Biogeochemical processes and chemical
behaviors of selenium
Speciation and bioavailability of selenium in soil
The pedosphere interacts directly with the lithosphere, the
hydrosphere, the atmosphere, and the biosphere, and plays
important roles in Se biogeochemistry cycling (Floor and
Román-Ross 2012). Adsorption, precipitation, and com-
plexation reactions can keep Se in soil, while desorption,
leaching, dissolution, and volatilization can remove Se from
the soil (Fig. 1) (Dinh et al. 2019). The transport and fate
of Se in soil can be affected significantly by soil proper-
ties particularly pH, Eh, and organic matter content (Cop-
pin et al. 2006; Li et al. 2016). Soil redox potential and pH
primarily determine soil Se speciation, and the chemical
transformation of Se would further alternate Se chemical
behaviors and regulate Se transport and fate in the soil,
which involves chemical and physical processes for adsorp-
tion and desorption between Se and soil components (Dinh
et al. 2019). As one of the oxidized Se chemical forms,
13
International Journal of Environmental Science and Technology
Location References
Armstrong et al. (2018)
Porth Dinllaen
Porth Dinllaen
Lianddwyn
Caemes Bay
Shotton and Northumberland Bullock et al. (2018)
selenite mainly exists in soil with moderate oxidation con-
ditions (pE + pH = 7.5–15.0), whereas selenate predominates
in high oxidizing conditions (pE + pH > 15.0) (Shahid et al.
2018). Selenate is water-soluble, with only a small portion
adsorbed to soil clay minerals. On the contrary, selenite can
be strongly fixed by minerals, metal hydroxides, and organic
matter in soils (El-Ramady et al. 2014; Zhai et al. 2019).
When soil pH was reduced, selenate could be transformed
to selenite, which particularly enhances Se adsorption with
metal (hydro)oxides and decreases Se bioavailability in
the soil (Dinh et al. 2019; Lee et al. 2011; Li et al. 2016;
Masscheleyn et al. 1990; Wang et al. 2017c). The adsorbed
selenite retained in the soil solid phase could be incorpo-
rated into the mineral structure. It may be possible to fur-
ther reduce it to metal selenides (such as Fe selenide) and
elemental Se (Borsig et al. 2017; Francisco et al. 2018; He
et al. 2018b; Wang et al. 2020b; Xu and Huang 2019). Mean-
while, some soil chemical compounds like ­MnO2 can also
oxidize selenite to selenate (Rosenfeld et al. 2020). When pH
increases, Se anions are released by electrostatic repulsion
from the soil (Dinh et al. 2019).
Selenium is present in soil in different components
according to the type of chemical form. Based on the
sequential extraction program, Se in soil can be classified
into five categories: soluble Se (SOL-Se), exchangeable Se
and carbonate-bound Se (EXC-Se), Fe/Mn oxide-bound Se
(FMO-Se), organic matter-bound Se (OM-Se), and residual
Se (RES-Se) (Wang et al. 2012). Among all components,
SOL-Se is the fraction most easily used by plants, while
EXC-Se can be desorbed from the soil and then assimilated
by plants (Wang et al. 2017b). Inorganic and organic Se
compounds adsorbing to soil organic matter are included
in the OM-Se fraction (Abrams et al. 1990). Soil organic
matter serves as a major carrier of Se in soil and the organic
acids within the organic matter has a crucial impact on Se
bioavailability (Wang et al. 2019; Qin et al. 2012). Soil
organic acids include low molecular weight organic acids
of succinic, malic, citric, and oxalic acid, as well as high
International Journal of Environmental Science and Technology
Table 2  Selenium contents in soils
Country/Region Se contents
America 3.75–435 mg ­Kg−1
6.94 mg ­Kg−1
2–5 mg ­Kg−1
0.6–26 mg ­Kg−1
1–3 mg ­Kg−1
10–12 mg ­Kg−1
0.4–20 mg ­Kg−1
China 5.15–63 mg ­Kg−1
0.039–1.110 mg ­Kg−1
0.03–16.96 mg ­Kg−1
0.03–86.59 mg ­Kg−1
0.17–2.89 mg ­Kg−1
0.11–1.48 mg ­Kg−1
61.8–462.6 μg ­Kg−1
0.02–4.68 mg ­Kg−1
0.01–4.26 mg ­Kg−1
France 25–1222 μg ­Kg−1
198–815 μg ­Kg−1
323–1119 μg ­Kg−1
172–1041 μg ­Kg−1
Korea 1.7–10.2 mg ­Kg−1
0.7–3.5 mg ­Kg−1
1.3–9 mg ­Kg−1
0.5–2.4 mg ­Kg−1
1.1–2.1 mg ­Kg−1
India 13.1 mg ­Kg−1
6.8 mg ­Kg−1
0.023–4.91 mg ­Kg−1
Ireland 13.5–105 mg ­Kg−1
0.6–175 mg ­Kg−1
217.3–1200 mg ­Kg−1
0.1–7.8 mg ­Kg−1
Nigeria 1.2–9.6 mg ­Kg−1
Poland < 0.3–0.8 mg K­ g−1
Saudi Arabia 0.1–0.7 mg ­Kg−1
Scottish 0.19–1.46 mg ­Kg−1
Slovakia 0.44–4.25 mg ­Kg−1
Norway 3–6 mg ­Kg−1
Scotland 0.115–0.877 mg ­Kg−1
Finland 0.005–1.241 mg ­Kg−1
New Zealand 0.1–4 mg ­Kg−1
England < 0.01–16 mg K ­ g−1
Germany 9.60–12.31 mg ­Kg−1
Egypt 17.78–42.70 mg ­Kg−1
molecular weight organic acids like humic acid and fulvic
acids (Adeleke et al. 2017). Both of them will contend for
adsorption sites with Se, which will further weaken soil Se
Location References
Western Phosphate Resource Area Favorito et al. (2017)
Western Colorado Statwick and Sher (2017)
Hawaii Byers et al. (1936)
Maui
Molokai
Oahu
Kauia
Enshi Qin et al. (2017)
Jiangjin Liu et al. (2021)
Langao Hao et al. (2021)
Enshi Li et al. (2020)
Guiyang (0–20 cm) Pan et al. (2017)
Guiyang (150–200 cm)
Luolong Wang et al. (2020c)
Basu
Hainan Island (0–20 cm) Gong et al. (2022)
Hainan Island (150–180 cm)
Forest Pisarek et al. (2021)
Agriculture Tolu et al. (2014)
Meadow
Forest
Dukpyung mountain Park et al. (2010)
Dukpyung farmland
Chubu mountain
Chubu farmland
Chubu paddy
Punjab Wheat soil Eiche et al. (2015)
Punjab Mustard soil
Punjab Dhillon and Dhillon (2014)
Limerick Fleming and Walsh (1957)
Tipperary
Meath
Northern Ireland (5–20 cm) Jackson et al. (2016)
Lower Benue trough and the Niger Delta Basin Nganje et al. (2020)
Holy Cross Mountains Galuszka (2005)
Farming areas located along the Gulf of Aqaba coast Ghrefat et al. (2016)
Topsoil Shand et al. (2010)
Surface soil (Sobov surface mine) Bujdos et al. (2005)
Fordyce (2013)
Wales
Floodplain soils along the Wupper River Shaheen et al. (2017)
Floodplain soils in the north of the Nile River delta
adsorption by metal hydroxides (Dynes and Huang 1997;
Fang et al. 2019; Masset et al. 2000; Zhou et al. 2007). Low
molecular weight organic acids can break the binding of
13
 International Journal of Environmental Science and Technology

Table 3  Selenium concentrations in waters

Country/Region Total dissolved Dissolved inor- Dissolved Se(IV) (μg ­L−1) Se(VI) (μg ­L−1) Types of water References
Se ganic Se organic Se
(μg ­L−1)

America < 0.5–4070 μg Groundwater Mills et al. (2016)

­L−1
1.1–17 μg ­L−1 Groundwater Beisner et al.
(2020)
0.017–4.46 μg 0–1.07 0–2.08 0–1.29 Groundwater Basu et al. (2007)
­L−1
0.02–0.069 μg 0–0.021 0–0.023 0–0.033 Groundwater
­L−1
Britain 0.31–2.83 μg Rainwater Blazina et al.
­L−1 (2017)
Canada 0.11–0.16 μg Athabasca River Donner et al.
­L−1 (2018)
China 0.004–0.09 0.0007–0.094 Changjiang Wu et al. (2014)
Estuary and
its adjacent
waters
0.08–0.55 μg 0.03–0.13 0–0.13 0.011–0.38 Changjiang Chang et al.
­L−1 Estuary (2021)
0.03–0.59 μg Inland rain Wang and Gao
­L−1 (2001)
0.015–0.11 0.008–0.099 Bohai Duan et al. (2010)
Chile 2.1–797 μg ­L−1 Groundwater Leybourne and
Cameron (2008)
France 0.013–0.18 μg < 0.006–0.015 0.011–0.067 Rainwater Suess et al. (2019)
­L−1
India 0.01–35.6 μg Groundwater Dhillon and
­L−1 Dhillon (2016)
2.5–69.5 μg ­L−1 Groundwater Dhillon and
Dhillon (2014)
Indonesia < 0.1–79 μg ­L−1 Groundwater Winkel et al.
(2008)
Ireland 0.01–5.63 μg Stream Waters Jackson et al.
­L−1 (2016)
Malaysia 0.055–0.45 μg 0–0.037 μg ­L−1 0.033–0.45 Rivers and Chang et al.
­L−1 estuaries (2020)
Norway < 0.5–1.4 μg Stream and river Reimann et al.
­L−1 water (2009)
Ocean 0.020 μg ­L−1 0.004 0.008 0.008 Surface ocean Mason et al.
(2018)
0.026 μg ­L−1 0.0008 0.016 0.010 Subsurface
0.11 μg ­L−1 0.045 0.068 Deep ocean
Pacific 0.038–0.061 μg North and South Mason et al.
­L−1 Pacific surface (2018)
water (< 10 m)
0.17 μg ­L−1 North and South
Pacific deep
water
(> 3000 m)
Pakistan 0.86–131.31 μg Groundwater Khan et al. (2010)
­L−1
Saudi Arabia 2.12–152.84 μg Groundwater Alqahtani et al.
­L−1 (2020)
0.1–3 μg ­L−1 Groundwater Ghrefat et al.
(2016)

13
International Journal of Environmental Science and Technology

Table 3  (continued)
Country/Region Total dissolved Dissolved inor- Dissolved Se(IV) (μg ­L−1) Se(VI) (μg ­L−1) Types of water References
Se ganic Se organic Se
(μg ­L−1)

Slovakia 0.12–32.18 μg Surface waters Bujdos et al.

­L−1 (2005)
Swiss 0.005–0.079 μg < 0.006–0.012 < 0.005–0.033 Alps rainwater Suess et al. (2019)
­L−1
Tanzania < 0.01–3.0 μg Tap water Tomasek et al.
­L−1 (2022)
0.3–7.1 μg ­L−1 Groundwater

Fig. 1  Schematic biogeochemical behavior of selenium in ecosys- thionine (SeMet), dimethyl selenide (DMSe), and dimethyl diselenide
tem. The selenium migration (white), oxidation (red), and reduction (DMDSe). Data of recommended nutrient intakes (RNI) and upper
(green) processes are presented by the corresponding arrows. The intake levels (UL) of selenium are from FAO/WHO, where F is for
organic selenium species include selenocystine (SeCys), selenome- female and M is for male

Se to soil materials, while organic acids with high molecu-
lar weight can be fixed by complexing or chelating with Se
(Dinh et al. 2017). The FMO-Se and RES-Se bind strongly
to components of the soil, such as metal oxides, and are
therefore poorly absorbed by plants. Meanwhile, soil micro-
organisms can absorb and assimilate Se into organic-Se like
selenoamino acids or produce Se nanoparticles (SeNPs)
through biological reduction (Fig. 1) (Zhu et al. 2018).
Emission and deposition of selenium in atmosphere
The transport and fate of Se in the atmosphere have sig-
nificant impacts on Se levels in soil and water, which ulti-
mately affects Se accumulation in crops. The atmosphere
is an important part of Se biogeochemical cycling. There
are approximately 29,000–36,000 tons of Se circulating
in the atmosphere annually, and the average life-time of

13
 International Journal of Environmental Science and Technology

Table 4  Selenium Country/Region ­ −3)

Se contents (ng m Location References
concentrations in the
atmosphere Baltic sea 0.1–1.3 Hoburg Dudzinska-Huczuk and Bolalek (2007)
0.14–2.33 Kap Arkona
0.08–1.69 Preila
China 4.1a Beijing Cui et al. (2020)
1.23–8.41a Nanjing Qi et al. (2016a)
0.8–5.0a Nanjing Qi et al. (2016b)
0.01–2.76b Shanghai Lv et al. (2012)
7.22–22.41 Xiamen Lee et al. (2009)
4.68–5.56c Guangzhou Xiao et al. (2014)
4.02–4.31a
Italy 2.56–284.69d Venice Rampazzo et al. (2008)
Polish 12.7a Warsaw Majewski and Rogula-Kozowska (2016)
Spain 0.2–2.5b Seville Enamorado-Baez et al. (2015)
Turkey 0.1–3.1d Düzce Urban Bozkurt et al. (2018)
0.1–1.7d Düzce Suburban

a: fine particles matter (­ PM2.5) b:particles c:total suspended particles d:coarse particles matter (­ PM10)

atmospheric Se is 4.4 days (Feinberg et al. 2020a, b). The
source emission to the atmosphere, long-range transport,
and atmospheric deposition of Se are important biogeo-
chemical processes that determine Se distribution and fate
in the environment (Feinberg et al. 2020a, b; Lanceleur
et al. 2019; Wen and Carignan 2007). Airborne Se comes
from both anthropogenic and natural. Natural Se emission
includes biogenic volatilization from marine, terrestrial
ecosystems, and volcanic emissions, and anthropogenic
emission sources include fossil fuel combustion and metal
smelting processes (Fig. 1). Overall, biogenic volatile Se
produced in the marine biosphere is the predominant natu-
ral Se source in the atmosphere, accounting for 39% of the
total Se emissions, compared to the terrestrial emission
sources of 15%. Volcanic eruption contributes about 12%
of the total Se in the atmosphere. The remaining 34% of
Se emissions are related to human activities such as coal
combustion, metal smelting, and biomass combustion
(Feinberg et al. 2020b). Volatile Se compounds emitted
from marine and terrestrial ecosystems are mainly dime-
thyl selenide (DMSe) and dimethyl diselenide (DMDSe)
formed by biological methylation (Fig. 1). Dimethyl sele-
nide is the predominant type of Se volatilization in terres-
trial ecosystems (Frankenberger and Karlson 2001), and
DMDSe is not stable in the environment and can easily be
transformed into ­Se0, organic Se and soil OM-Se (Zhang
and Frankenberger 2002). Volatile Se species can also be
oxidized to inorganic species within minutes to hours after
they are released into the atmosphere. Dimethyl selenide
will react with ­O2, ­H2O2, OH, ­NO3, and other substances
to form DMSeO that can be further converted to inorganic
Se through a series of oxidation reactions (Fig. 1) (Wen
and Carignan 2007).
CH3 SeCH3 + 2H2 O2 → CH3 SeO2 + 2H2 O
CH3 SeCH3 + OH → CH3 SeCH⋅2 + H2 O
( )
CH3 SeCH3 + NO3 → CH3 Se ONO2 CH3 → CH3 SeCH⋅2 + HNO3
CH3 SeCH⋅2 + NO−3 → CH3 SeCH2 O (DMSeO) + NO−2
2CH3 SeCH⋅2 + O2 → 2CH3 SeCH2 O
Volcanoes and human activities mainly release inorganic
Se such as Se dioxide (­ SeO2), elemental Se (­ Se0), and hydro-
gen selenide (­ H2Se) into the atmosphere (Fig. 1). It could
be expected that ­H2Se could be oxidized to ­SeO2 that can
dissolve in rainwater and form H ­ 2SeO3 (Fig. 1). Meanwhile,
­SeO2 can also react with ­SO2 in water to form ­Se0. The ­Se0
and ­SeO2 generated from these reactions can also be con-
verted into particulate Se, which can be long-range trans-
ported in the atmosphere (Floor and Román-Ross 2012). The
reaction between S­ e0 and sulfoxy radicals transforms insolu-
0
ble ­Se to selenate and selenite (Bronikowski et al. 2000;
Floor and Román-Ross 2012; Wen and Carignan 2007).
SeO2 + H2 O = H2 SeO3
H2 SeO3 + 2SO2 + H2 O = Se0 + 2H2 SO4
SeO2 + 2SO2 = Se0 + 2SO3
Lee et al. (2015) found that concentrations of Se in
Greenland snow were affected by coal combustion through

13
International Journal of Environmental Science and Technology
long-distance transport. Dry and wet deposition are the
means by which atmospheric Se eventually returns to the
Earth surface, in which wet deposition accounts for about
80% of global total Se deposition (Fig. 1) (Feinberg et al.
2020a, b). Haygarth et al. (1995) studied the effects of
atmospheric Se on plants by isotope dilution. The results
showed that nearly half (53%) of the Se in pasture leaves
came from the atmosphere. This research suggests that the
atmosphere is also a source of Se for plant uptake and can
affect human Se dietary intake through the food chain. It is
noteworthy that many atmospheric transformations of Se are
still unclear (Wen and Carignan 2007).
Migration and transformation of selenium in water
In aquatic environments, selenate and selenite are two
dominant chemical forms. Redox potential and pH value
have significant impacts on water Se speciation. Selenate
mainly exists under strong aerobic and oxidizing condi-
tions, and selenite is the predominant Se form in a slightly
reducing environment, mostly as monohydrogenium selenite
­(HSeO3−) in natural waters (El-Ramady et al. 2014; Leblanc
et al. 2018). In seawater, the contents of Se increase with
ocean depth increasing. Few studies have been conducted
to determine Se speciation in seafood materials under ocean
environmental conditions. It was expected that organic Se
presents primarily in the photic zone (or the upper mixed
layer) as the results of Se assimilation and biotransforma-
tion by bacteria and phytoplankton, meanwhile, there is a
vital relationship between Se and phytoplankton productiv-
ity (Fig. 1) (Duan et al. 2010). Aquatic organisms absorb and
transform inorganic Se into organic forms and further vola-
tile Se compounds through biological methylation (Fig. 1).
The decomposition remnants of aquatic organisms are
sunken to the water bottom and carrying Se to the sediment
(Fig. 1) (Graves et al. 2019; Zhou et al. 2019). The major
forms of Se in sediments are organic Se and elemental Se
­(Se0) (Janz et al. 2014). Elemental Se formation could be
enhanced through the buildup of detrital materials that sup-
port higher microbial activities and biological reduction of
inorganic Se at the surface of sediments (Fig. 1) (Zhao et al.
2020; Zhou et al. 2019). The species analysis of organic Se
in sediments showed that the main organic Se was SeMet,
which was then transformed into volatile Se and left into
the water body (Zhao et al. 2020; Zhou et al. 2019). Mean-
while, earlier research also demonstrated that elemental Se
in freshwater sediments could also be oxidized into selenite
at the water–sediment interface and selenite could be further
absorbed by organisms (Fig. 1) (Janz et al. 2014). The Se
transport through detrital and benthic food chains in aquatic
environments has been previously addressed (Graves et al.
2019; Janz et al. 2014; Zhou et al. 2019). The boundary
transport of Se between water and sediment has a significant
role in Se biogeochemical cycling and significantly influ-
ences the distribution of Se in surface water. While surface
runoff from Se-rich soils will bring Se to surface water, the
Se content in groundwater can be increased through soil Se
leaching from excessive irrigation and rainfall, along with Se
released from rock weathering (Kumar et al. 2011).
For the human Se intake, drinking water and other bever-
ages contribute between 5 and 25% of the Se intake (Kiel-
iszek 2019). A significant source of dietary Se is fish, whose
Se content is influenced by Se levels in water (Graves et al.
2019). Amongst different countries, fish contribute 60%,
21%, 13%, 9%, and 8% of the daily Se dietary intake in
Japan, Korea, UK, Brazil, and Argentina, respectively (Choi
et al. 2009; Haratake et al. 2007; Retondario et al. 2019;
Sigrist et al. 2012; Ysart et al. 2000). Concentrations of Se
are generally high in fish caught from the high-latitudes
ocean (Hicks et al. 2019). For instance, the Se concentration
in fish caught from North Korea (0.359 μg ­g−1), Argentina
(0.315 μg ­g−1), and Greenland (0.268 μg ­g−1) was higher
compared to that in fish caught from the low-latitudes like
Poland (0.192 μg ­g−1), Germany (0.182 μg ­g−1), and Malta
(0.175 μg ­g−1) (Hicks et al. 2019).
Biological accumulation and transformation
of selenium
Activity and toxicity of Se in organisms depend on Se spe-
cies and quantity. Compared with selenate and selenite, sele-
noamino acids, like ­SeCys2, MeSeCys, and SeMet are more
easily absorbed and less toxic to organisms (Rayman 2020).
Nanoscale elemental Se particles have recently gained much
attention for their multiple properties including biocompat-
ibility, bioavailability, low toxicity, and antimicrobial prop-
erties. Among inorganic Se in the environment, selenate and
selenite are predominant, and their reduction to selenoamino
acid and SeNPs is mainly driven by plants, algae, and micro-
organisms (Fig. 1).
Microorganisms and plants can biotransform of Se through
oxidation, reduction and methylation (Fig. 1) (Nancharaiah
and Lens 2015; Praus and Szakova 2019). Plants can absorb
selenate, selenite, organic Se, and SeNPs in the soil (Fig. 1)
(White 2016). Selenate can be transported from roots to leaves
through the xylem system for assimilation, while selenite will
be oxidized into selenate or reduced into organic selenide in
plant roots (Hu et al. 2018; Wang et al. 2015; White 2018).
The plant Se assimilation pathway shows that selenate is first
converted to adenosine 5’-phosphoselenate (APSe) by aden-
osine triphosphate sulfurylase (ATPS), and then reduced to
selenite by adenosine 5’-phosphosulphate reductase (APR)
with reduced glutathione (GSH) as electron donor (White
2016, 2018). Selenite can be converted into selenides by
intracellular sulfite reductase (SiR) or reduced glutathione,
and subsequently, selenide reacts with O-acetylserine (OAS)
13

to form SeCys with cysteine synthase (Gupta and Gupta 2017;
Schiavon and Pilon-Smits 2017; Wirtz and Hell 2006). Sele-
nomethionine is synthesized from SeCys, which is first con-
verted to selenocystathionine (SeCysTH) by cystathionine
γ-synthase (CGS), then to selenohomocysteine (SeHCys) by
cystathionine β-lyase (CBL), and then to SeMet by methio-
nine synthase (MTR) (Trippe and Pilon-Smits 2021; White
2018). These selenoamino acids can be involved in protein
synthesis in plants, which plays a significant physiological
function in producing volatile Se compounds like DMSe and
DMDSe through biomethylation (Fig. 1) (Kolbert et al. 2019;
Schiavon and Pilon-Smits 2017; Tagmount et al. 2002). Algae
can absorb inorganic Se from water and reduce to selenoamino
acids via the sulfur reduction pathway that is similar to the Se
metabolism pathway in plants (Schiavon et al. 2017; Vriens
et al. 2016).
Bacteria and fungi can also absorb Se from the environ-
ment, which is the first step in microbial Se metabolism. The
absorption of selenate occurs mainly via the sulfate trans-
porter, whereas the absorption of selenite is associated with
sulfate and phosphate transporter systems (Avendano et al.
2023; Tan et al. 2018; Turner et al. 1998; Zhu et al. 2020).
It has been shown that the polyol transporter, the tripartite
ATP-independent periplasmic (TRAP) transporter TakP, and
the membrane protein DedA are also involved in the uptake
of selenite by bacteria (Adnan et al. 2021; Bebien et al. 2001;
Ledgham et al. 2005). Meanwhile, in Rahnella aquatilis
HX2, selenite transportation is mediated by active trans-
port processes and water channel protein (AqpZ) (Xu et al.
2023b). For bacteria, Se reduction ability have been identi-
fied in the phylum of Proteobacteria, Firmicutes, and Act-
inobacteria (Siddique et al. 2005). Microorganisms are able
to use selenate as a terminal electron acceptor for anaerobic
respiration. When selenate enters cells, it is reduced to selenite
under the action of selenate reductase and then reduced to
insoluble ­Se0. Selenate reductase was identified in Thauera
selenatis, Enterobacter cloacae, Bacillus selenatarsenatis,
Escherichia coli, Salmonella enterica, and Citrobacter fre-
undii (Butler et al. 2012; Guymer et al. 2009; Kuroda et al.
2011; Ridley et al. 2006; Theisen and Yee 2014). In anaero-
bic conditions, selenate reductases can reduce selenate in the
periplasm with follow process (Debieux et al. 2011; Guymer
et al. 2009; Kuroda et al. 2011; Ridley et al. 2006; Schroder
et al. 1997). However, in aerobic conditions, the reduction of
selenate mainly via the sulfate reduction pathway (Tan et al.
2018). In fungi, the reduce mechanism of selenate is through
the sulfur reduction pathway, with ATP sulfurylase and APSe
kinase (Kieliszek et al. 2015).
SeO2−
4
+ 2e− + 2H+ → SeO2−
3
+ H2 O
SeO2− + 4e− + 6H+ → Se0 + 3H2 O
3
13
International Journal of Environmental Science and Technology
In bacteria, selenite reduction occurs in the periplasm
or cytoplasm. Some research showed that selenite reduc-
tase also exists in microbes. Wells et al. (2019) identified a
respiratory selenite reductase in Bacillus selenitireducens,
and CsrF was also confirmed as bacterial aerobic selenite
reductase in Alishewanella (Xia et al. 2018). In Comamonas
testosteroni S44, a periplasm molybdenum oxidoreductase
SerT plays an important role in the selenite reduction pro-
cess (Tan et al. 2018), and the selenite reductase FesR can
reduce chromate in Alishewanella sp. WH16-1 (Zhou et al.
2022). Meanwhile, some studies have shown that nitrite
reductase, fumarate reductase, hydrogenase I, and sulfite
reductase are involved in the selenite reduction process
(Demolldecker and Macy 1993; Huang et al. 2021; Jia et al.
2022; Li et al. 2015; Yanke et al. 1995). It is evident that the
selenite reduction process in microorganisms involves mul-
tiple pathways, including Painter-type reaction, thioredoxin-
thioredoxin reductase system, siderophore-mediated reduc-
tion, sulfide-mediated reduction, and dissimilatory reduction
(Nancharaiah and Lens 2015). The Painter-type reduction
mediated by glutathione (GSH) and its homologs may be the
primary process involved in selenite reduction (Nancharaiah
and Lens 2015; Wang et al. 2022a). In the reaction, selenite
first reacts with glutathione to form selenodiglutathione
(GSSeSG), which is subsequently converted to selenop-
ersulfide anion (GS-Se−) under the action of glutathione
reductase (Ganther 1971; Kessi and Hanselmann 2004).
After undergoing hydrolysis reaction, the GS-Se− decom-
poses to form S ­ e0, which will subsequently assembled into
SeNPs (Ganther 1971; Tugarova and Kamnev 2017).
6GSH + 3SeO2−
3
→ 3GS − Se − SG + 3O−2 + 3H2 O
GS − Se − SG + NADPH → GSH + GS − Se− + NADP+
GS − Se− + H+ → GSH + Se0
The reduction of selenite by bacteria has been influenced
by other factors. The small RNA chaperone Hfq regulated
bacterial selenite reduction and SeNPs biosynthesis by sul-
fate and glutathione reduction systems in Rahnella aqua-
tilis (Xu et al. 2020). These biosynthetic SeNPs have been
observed in inside and outside of bacterial cells (Zhu et al.
2018), which are assembled via flagellin FliC and porin
OmpF in R. aquatilis (Li et al. 2021). However, the secre-
tion channels of SeNPs is currently unclear. Microorgan-
isms can assimilate inorganic Se into selenoamino acids
(SeCys and SeMet) for accumulation in the body and can
also be used to synthesize selenoproteins (Hu et al. 2019a,
b, 2020, 2021; Martinez et al. 2020; Zhao et al. 2023). At
the cellular level, methyltransferases catalyze the transfer
of methyl from organic substrates, which produce volatile
methylated products as DMSe and DMDSe (Winkel et al.
International Journal of Environmental Science and Technology
2015). Previous studies have found that the bacterial thio-
purine methyltransferase (bTPMT) plays an important role
in the Se methylation process (Ranjard et al. 2002, 2003). In
contrast, the fungus synthesizes volatile Se via 5-methyltet-
rahydropteroyltriglutamate-homocysteine methyltransferase
(MetE) (Xu et al. 2023c). The biological methylation and its
volatilization to the atmosphere is an important process in Se
biogeochemical cycling, which enhance the mobility of Se in
the environment and lead to redistribution of Se in the envi-
ronment (Winkel et al. 2015). In addition, microorganisms
can also oxidise ­Se0 and organic Se (SeMet, ­SeCys2, and
selenourea) to selenate and selenite, and the rate constant of
oxidation is much lower than that of reduction (Dowdle and
Oremland 1998; Luo et al. 2022; Zhu et al. 2021).
The biogeochemical cycling of Se in the terrestrial eco-
system involves Se uptake by primary producers, assimi-
lated by herbivores and then carnivores, with possible bio-
magnification through the trophic levels (Fig. 1) (Delariva
et al. 2017; Trumble et al. 1998; Vickerman and Trumbleet
2003). It was suggested that biomagnification of Se would
also increase Se bioavailability to organisms at higher
trophic levels (Hopkins et al. 2005). Selenium accumulated
in insects mainly exists as organic forms and becomes an
important food source for other predators (Fig. 1) (Andra-
hennadi et al. 2007; Golubkina et al. 2014). Moreover, soil
animals can also directly absorb Se and convert inorganic Se
into organic Se (Azhar-u-ddin et al. 2020) as major consum-
ers and important food sources for many organisms (Andra-
hennadi et al. 2007; Golubkina et al. 2014). The decomposi-
tion of soil animal and plant tissues will also directly result
in the addition of Se to the soil (Quinn et al. 2010). The
absorption and transformation of Se by different organisms
will be important steps affecting the biogeochemical cycling
process of Se and eventually affect human health through the
food chain (Fig. 1).
Conclusion
Microbes including bacteria and fungi play significant roles
in the biogeochemical processes of Se in both terrestrial and
aquatic ecosystems. Plants, bacteria, fungi, and algae can
absorb and reduce selenate and/or selenite to selenoamino
acids, SeNPs, and/or methyl Se, and play significant roles
in the biogeochemical processes of Se in both terrestrial
and aquatic ecosystems. Selenium content and distribution
in soil are significantly affected by Se biogeochemical pro-
cesses that will affect human Se dietary intake. Clearly, it is
important to better manipulate natural Se biogeochemical
processes in order to effectively manage human Se dietary
intake based on the RDA in Se deficiency regions. There are
several aspects that should be emphasized.
(1) The transformation process of Se in different layers
of the Earth has been thoroughly studied. To further
understand the cycle of Se, it is requisite to further
explore the geochemical behavior of Se, especially the
transfer and transformation process of Se at two differ-
ent environmental interfaces.
(2) Microorganisms are crucial to the geochemical cycle
of Se, and the absorption and transformation of Se by
bacteria and fungi have been thoroughly studied. As
the third domain of life, archaea are widely distributed
in the environment. However, there are few studies on
the biotransformation of Se by archaea, which should
be paid attention to in future studies.
(3) The biogeochemical processes of Se can significantly
affect its distribution in the environment and thus affect
the human Se intake. Therefore, it is worth exploring
whether Se distribution in the environment can be
improved by regulating the cycling process of Se to
improve human health.
Author contribution YG conceived and designed the manuscript; ZX,
ZL, and YG wrote the manuscript; ZX collected literature and data;
GZ revised the manuscript. All authors have read and agreed to the
published version of the manuscript.
Funding This work was supported by Special Fund for Key Science &
Technology Program in Xinjiang Province of China (No. 2022B02053
& 2022B02021) and Special Fund for Agro-Scientific Research in the
Public Interest of China (201303106).
Data availability Not applicable.
Declarations
Conflict of interest The authors declare no competing interests.
Ethical approval Not applicable.
Consent for publication Not applicable.
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