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Fuentes-Lara, L.O.; Medrano-Macías, J.; Pérez-Labrada, F.; Rivas-Martínez, E.N.; García-Enciso, E.L.; González-Morales, S.; Juárez-Maldonado, A.; Rincón-Sánchez, F.; Benavides-Mendoza, A. Transformations of Elemental Sulfur in Soil. Encyclopedia. Available online: https://encyclopedia.pub/entry/51777 (accessed on 17 May 2024).
Fuentes-Lara LO, Medrano-Macías J, Pérez-Labrada F, Rivas-Martínez EN, García-Enciso EL, González-Morales S, et al. Transformations of Elemental Sulfur in Soil. Encyclopedia. Available at: https://encyclopedia.pub/entry/51777. Accessed May 17, 2024.
Fuentes-Lara, Laura Olivia, Julia Medrano-Macías, Fabián Pérez-Labrada, Erika Nohemí Rivas-Martínez, Ema Laura García-Enciso, Susana González-Morales, Antonio Juárez-Maldonado, Froylán Rincón-Sánchez, Adalberto Benavides-Mendoza. "Transformations of Elemental Sulfur in Soil" Encyclopedia, https://encyclopedia.pub/entry/51777 (accessed May 17, 2024).
Fuentes-Lara, L.O., Medrano-Macías, J., Pérez-Labrada, F., Rivas-Martínez, E.N., García-Enciso, E.L., González-Morales, S., Juárez-Maldonado, A., Rincón-Sánchez, F., & Benavides-Mendoza, A. (2023, November 18). Transformations of Elemental Sulfur in Soil. In Encyclopedia. https://encyclopedia.pub/entry/51777
Fuentes-Lara, Laura Olivia, et al. "Transformations of Elemental Sulfur in Soil." Encyclopedia. Web. 18 November, 2023.
Transformations of Elemental Sulfur in Soil
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This entry is adapted from the peer-reviewed paper 10.3390/molecules24122282

Sulfur is an essential element in determining the productivity and quality of agricultural products. It is also an element associated with tolerance to biotic and abiotic stress in plants. In agricultural practice, sulfur has broad use in the form of sulfate fertilizers and, to a lesser extent, as sulfite biostimulants.

sulfate sulfite nutraceuticals polysulfanes polysulfides soil microbiome

1. Introduction

Sulfur is one of the most abundant elements on Earth and is an essential element for living beings, of which constitutes on average 1% of dry weight. In plants, S content varies strongly between species, ranging from 0.1 to 6% of dry weight (0.03 to 2 mmol g−1 dry weight) [1]. S belongs to the VIA group of the periodic system, where it is found together with O, Se, Te, and Po; naturally, S is a mixture of four isotopes, 32S, 33S, 34S, and 35S. The natural abundance of each is 95.1%, 0.74%, 4.2%, and 0.016%, respectively. Sulfur exists in oxidation states ranging from +6 to −2 (Table 1), with the most oxidized state in the form of sulfate (SO42−), which is the chemical form that plants absorb from the soil to feed themselves with S [2].
Table 1. Representative sulfur compounds and their oxidation states.

Oxidation State

Representative Compound and Formula

Oxidation State

Representative Compound and Formula

+6

Sulfate, SO42−

0

S0, elemental sulfur. Sulfoxide (R-S(-O)-R such as dimethyl sulfoxide (DMSO). Oxidized derivatives of sulfide and sulfenic acid (RSOH).

+6 and −2

Thiosulfate, S2O32−

−1

Disulfide (R-S-S-R) is a persulfide found in the linkages between two cysteine residues in proteins. RSSH denotes persulfides (or hydrosulfides) obtained by the action of H2S on cysteine residues (R-SH). Thioethers and thiols can be oxidized to disulfides. Major products of decomposition of persulfides are polysulfanes. Thiyl-radical RS*.

+5 and −2

Polythionates (−O3S-Sn-SO3−): Dithionate, S2O62−; Trithionate, S3O62−; Tetrathionate, S4O62−

−2

Sulfide, S2−, polysulfides, S22−, S32−, S52−; carbon disulfide (CS2); FeS2; NaHS and Na2S are sources of S2− and of its conjugated acids SH− and H2S. Polysulfides (with Sn > 2) contain S0 atoms, which allows a diversity of oxidation states.

+4

Sulfur dioxide, SO2; Sulfite, SO32−; Disulfite, S2O52−; Sulfone, OS(S) the oxidation product of sulfoxides

−2

Hydrogen sulfide (H2S), disulfane (H2S2), and polysulfanes (RSSnSR, n > 2). Polysulfanes contain S0 atoms, which allows a diversity of oxidation states.

+3

Dithionite, S2O42−

−2

Thioethers (C-S-C) such as dimethyl sulfide (DMS), CH3-S-CH3 and dimethyl disulfide (DMDS), CH3-S-S-CH3.

+2

Carbonyl sulfide (COS), OCS

−2

Thiols (R-SH) such as glutathione (GSH) and methyl mercaptan, CH3-SH. Thiols are derived from the sulfhydryl group -SH of cysteine, which enables multiple oxidation states (−2 to +6). Thiolates are derivatives of thiols in which a metal or other cation replaces H.

0

Elementary sulfur (S0), mainly S8 (cycloocta-S)

−2

Carbon disulfide, CS2.
Biological molecules, which range from small molecules to proteins and other polymers, contain S in its more reduced states 0, −1, and −2. For example, it is known that approximately 40% of enzymes depend for their catalytic activity on the presence of sulfhydryl groups (-SH). These -SH groups participate in redox reactions, provide binding sites for toxic or physiologically important metals, and are related to the detoxification of various xenobiotics. It is also known that the tertiary and quaternary structure of many proteins is the result of the presence of disulfane bonds (-S-S-) formed by the oxidation of -SH groups of cysteine, a sulfur amino acid that, together with methionine, is a key factor in determining the nutritional value of plants, as well as a central element in the metabolism of S in all organisms [2].
For the above reasons, a close relationship between nitrogen and sulfur nutritional status has been found in plants [3][4]. Approximately 80% of nitrogen and sulfur incorporated in organic compounds of plants is found in proteins when both elements are in adequate proportions. The S/N balance of a plant, described by the organic S/N ratio, is in the range of 0.025 (legumes) to 0.032 (grasses) and is relatively constant from one species to another. Therefore, the amount of S required by a plant is strongly dependent on its N nutrition. The consequence is that the availability of S below the needs of the crops does not allow the adequate use of applied N [5].
Compounds as important as β-lactam antibiotics (penicillins, cephalosporins, and cephamycins) have an S atom derived from cysteine. The sulfur compound S-adenosyl-l-methionine (SAM) is the most crucial methylating agent known in all organisms; SAM-mediated transmethylation reactions are essential in the regulation of gene expression, the activity of various enzymes, the synthesis of compounds such as the osmolyte DMSP (dimethyl sulfoniopropionate) and DMS (dimethyl sulfide) gas, as well as in the production of antibiotics [2].
The Earth’s S stores are located in the lithosphere, hydrosphere, atmosphere, and biosphere. Human activities result in the extraction of S from the lithosphere (burning of fossil fuels, mining of elemental S and metals) and biosphere (oxidation of organic matter from the soil and burning of biomass). Anthropogenic S is incorporated into the global cycle mainly in the form of SO2 emitted into the atmosphere [6].
Between the terrestrial and marine masses, there is a constant flow of S via the atmosphere through the gaseous forms of the element (SO2, COS, H2S, DMS, and CS2) [6] and aerosols (mainly SO42− from the oxidation of sulfur gases, and <10% of organosulfates) [7], or by runoff from terrestrial to oceanic regions (Figure 1). The constant mobilization of S causes changes in the sulfur species that move from one terrestrial compartment to another. Under oxic conditions, the predominant inorganic form of S is SO42−, resulting from atmospheric deposition or oxidation of reduced forms of S. In the soil, continuous land tillage that oxidizes soil organic matter and repeated extractions for crops cause the decrease of S stores; for this reason, the regular application of S with the fertilizers is recommended [8].
Figure 1. Simplified biogeochemical sulfur cycle. Human activities, fauna, vegetation, and soil microorganisms can be visualized as an interface (as source and sink) to accelerate the transfer of sulfur species between the lithosphere, atmosphere, and hydrosphere.
In soils, most S is found in organic forms; the inorganic forms are elemental sulfur (S0) or SO42−, the latter can be found as gypsum or be adsorbed in the inorganic exchange matrix. The SO42− adsorbed in the soil is in dynamic equilibrium with the soil solution, and the adsorption/desorption quotient inversely depends on the pH value of the soil and the cations present in the exchange matrix, showing higher affinity for Al3+ > Ca2+ > K+ [9][10].
In soil, SO42− is subject to dissimilatory and assimilatory reduction. Dissimilatory reduction occurs when SO42− is used as a final acceptor of electrons in the anaerobic metabolism of microorganisms, producing H2S that is reoxidized in the presence of O2 or volatilized into the atmosphere. Assimilatory reduction is used by prokaryotes, algae, plants, and fungi for the biosynthesis of organic compounds, e.g., amino acids. Animals and protists cannot perform assimilatory reduction of SO42−; therefore, they depend on the organic sulfur compounds synthesized by other organisms [6]. In many crop species, sulfur is an element associated with nutritional quality and density of mineral nutrients, tolerance to stress, and the management of certain pests and pathogens [11][12][13].
In agricultural soils, SO42− used by crop plants comes mainly from the contribution of fertilizers with sulfates, such as ammonium sulfate, gypsum, potassium sulfate, magnesium sulfate, single superphosphate, ammonium phosphate sulfate, potassium magnesium sulfate, and sulfates of micronutrients [8][14], as well as the oxidation of S0, and of S2− contained in organic fertilizers. Another part of the S of crops is obtained from SO42− and aerosols coming from precipitation, as well as the absorption by soil and plants of aerosols and gases such as H2S, COS, and DMS. When S is added to the soil in the form of SO42−, plants and aerobic prokaryotes absorb it and incorporate it into a reductive metabolism that produces sulfide (S2−). On the other hand, when S is supplied as S0 or in the form of organic fertilizers (S2−), it must be oxidized to SO42− by the action of soil prokaryotes to be available to plants [2][6][8].

2. Transformations of Elemental Sulfur in Soil

The S available for plants in agricultural ecosystems is in dynamic storage (Figure 2). It comes from gaseous forms and aerosols of S from the atmosphere, from dissolved S (mostly SO42−) in rain and snow precipitation, and from SO42−, which is obtained from the oxidation of S of soil organic matter and S0. Sulfates can be fixed in the soil exchange matrix or leached to the subsoil [10]. In arid regions, SO42− can be stored in large quantities as gypsum in the subsoil, but in areas with higher water availability, leached SO42− is mobilized to lower horizons and to the subsoil [15].
Figure 2. Schematic representation of the flow of sulfur in soil. APS = adenosine 5′-phosphosulfate. Oxidation states of sulfur in the different molecules are: SO42− (+6); S2O62− (+5 and −2); S4O62− (+5 and −2); S3O62− (+5 and −2); SO32− (+4); SO2 (+4); S2O32− (+6 and −2); COS (+2); S0 (0); SH− (−2); S2− (−2); DMS (−2); CS2 (−2).
In the anoxic zones of the soil, S0 and SO42− are transformed to H2S that is volatilized or is reoxidized to S0 y and sulfate in the oxic zone. Plants and microorganisms take the SO42− and reduce it to S2− to incorporate it into a huge variety of organic compounds. Subsequently, these same plants and microorganisms transform a part of the sulfur to H2S, DMS, and CS2 [8][16]. The above volatile molecules have been associated with detoxification metabolism, stress tolerance, and signaling in plants and prokaryotes [17][18]. As with iodine [19], soil organic matter can transform the S to volatile forms by means of abiotic reactions, but the rate of transformation is very low in comparison with biotic metabolism of S [16].
Since there are several access ways by which S can enter the agricultural ecosystem, it is not possible to mark a specific starting point. Therefore, arbitrarily, the assumption of an application of S0 to the soil is taken, and the transformations that this material experiences up to SO42− are described. Once in the form of SO42−, it is assimilated into plant cells in the form of myriad organic compounds. The final part of the flow of S from soil to plants ends with the production of volatile compounds by plant cells, or in the transformation of the S contained in plant waste (Figure 2).
S atoms tend to avoid double bonds, therefore, in the S0, instead of forming molecules of S2 (S=S) the S atoms are grouped in the form of cyclic allotropes (cyclosulfur) or as long chains Sn (catena sulfur) [20]. The S0 used to apply to soil consists mainly of molecules of S8 (cycloocta-S) that are grouped, forming polymers of variable size; S8 is the most stable form from a thermodynamic point of view. S8 is a very electrophilic Lewis acid, so it reacts with nucleophilic anions or Lewis bases such as OH−, sulfides (S2−), thiols (R-SH), thiolates (RS−), I−, CN−, and SO32− [21][22].
S0 is applied to the soil or substrate in quantities ranging from 20 to 250 kg ha−1 yr−1, the last figure being equivalent to 200 mg S0 kg−1 soil. Once in the soil or substrate, S0 begins to transform into other chemical forms, mainly through biotic processes, and, to a lesser extent, by abiotic processes. The transformation rate is inversely proportional to the particle size and directly proportional to the temperature (Q10 = 4.0), humidity availability, and abundance of edaphic microorganisms [8][23][24].
Any factor that decreases bacterial activity, such as temperatures <10 °C or >40 °C and lack of humidity in the soil, will reduce the transformation of S. Flooded or compact soils will have anoxic conditions that induce high rates of conversion of S0 and SO42− into gaseous forms of sulfur [8][24]. The metabolism of S in soils can modify other processes, as in rice paddies, where the use of gypsum amendment has been shown to decrease greenhouse methane emissions [25]. In alkaline soils, it has been observed that the use of S0 induces acidification (by H2SO4), which increases the bioavailability of elements such as P [26].
When it is desired that S0 produces SO42− rapidly available for crops, an S0 source with a small particle size (<150 μm or 100 mesh) should be chosen. Contrarily, if a long-term impact (two or more consecutive crops) is sought, it is desirable to use S0 sources with a larger particle diameter, or even granular forms such as S0 prills or S0-fortified N-P-K and DAP fertilizers [27][28]. At a temperature of 14 °C, it was found that, in 51 weeks, 51% of S0 with particle diameter 41 μm (300 mesh) was oxidized, compared to 18% of S0 with 125 μm (120 mesh). In soils with low temperatures, S0 sized 41 μm will oxidize at a rate equivalent to S0 sized 125 μm in soils with higher temperatures [23]. In another experiment, applying 50 kg ha−1 of S0, it was found that 80–90% of S0 with particles <150 μm was oxidized over a period of 340 days [29].
On the other hand, it has been found that repeated applications of S0 to soil increase the population and the activity of oxidizing bacteria of S0 [24]. Accompanying the increase in S0 oxidant bacteria was a reduction in the number of fungi and protists, while bacterial and actinomycete populations remained stable [30]. Other authors reported a decrease in biomass and bacterial metabolism by applying S0 annually for five years [31].
When S0 is in micronized form (<177 μm, <80 mesh) it is used for the control of mites and some fungi [32][33]. The reactivity of micronized S0 is a consequence of the high quotient surface/volume of the particles, estimated to be 1300 to 1940 cm2 g−1 for S0 of 125 and 41 μm, respectively [23]. Micronized S0 can be applied through the foliar route or even by using pressurized irrigation systems to incorporate it into the soil [34][35]. When applied by irrigation system, the problems associated with the application of micronized S0 (because it is a flammable and irritant material) by dusting machines are reduced [24].
The use of S nanoparticles for the control of pathogens in plants has also been described [36][37]. Taking into account the high value of the surface/volume ratio of S nanoparticles, furthermore being a source of S for rapid assimilation by plants and microorganisms, it is possible that they function as biostimulants [38], and that they provide highly reactive S0 that works as a tolerance-inducing factor against pathogenic fungi [32][39].
To be available for plants, S0 applied to the soil or substrate must be oxidized to SO42−. The change in the oxidation state of sulfur from 0 to +6 allows reduction equivalents to be obtained (8H+ + 6e−). The oxidation is carried out by most soil microorganisms, highlighting Thiobacillus, Beggiatoa, Desulfomicrobium, and Desulfovibrio, as well as other heterotrophic aerobics S-oxidizing bacteria such as Bacillus, Pseudomonas, and Arthrobacter [2][40]. Two metabolic pathways have been described that allow the oxidation of inorganic S to SO42−: the Kelly–Friedrich pathway, which does not involve the production of intermediates such as polythionates, and the Kelly–Trudinger pathway, which includes as an intermediate output tetrathionate (S4O62−) and other polythionates [16]. The existence of two different routes and the large number of taxa that carry out the oxidation of S0 allow a high redundancy, and capacity to tolerate extensive changes in pH and salinity in soils [41][42].
In Figure 2, the oxidation activity from S0 to SO42− is presented on the right side, and shows the Kelly–Trudinger pathway with the production of polythionates such as S4O62− and S3O62− (as well as S2O32− and SO32−), which serve as a source of reducing potential and possibly act as inducers of stress tolerance in plants, perhaps by containing a single sulfane sulfur not associated with oxygen [16].
Under anoxic conditions, S0 is produced as part of the dissimilatory reduction of SO42−. Later, the S0 can be assimilated into S2− that will be part of the biomolecules, or it will be volatilized in the case of excess S (see the central section of Figure 2). At the left and right ends of Figure 2, in the central part, the SO42− from fertilizers, precipitation, and mineralization of organic matter is represented. A portion of this SO42− forms a soil adsorbed sulfate storage, which will be in dynamic equilibrium with SO42− dissolved in the soil solution. Under oxic conditions, SO42− is assimilated in S2− by assimilatory reduction and then transformed back into SO42− during the organic decay and mineralization of organic matter [42].
As part of the processes of organic decay, mineralization of organic matter, and sulfate reduction, both the soil, through abiotic reactions, and micro-organisms and plants can be source or sink of volatile forms of S, such as H2S, DMS, COS, CS2, and SO2 (Figure 2). Generally, under anoxic conditions, the oxidized forms of S are reduced by the soil microbiome to H2S, CS2, COS, DMDS, methyl mercaptan, and COS [8]. These gaseous molecules are believed to be part of a mechanism of dissipation of excess S, although participation in other processes is not ruled out [16][43].
In terms of reductive and oxidative microbial reactions, the most abundant forms of sulfur in the soil and the edaphic microbiome are S2−, R-SH, RSSH, polysulfides (RSn2−), S2O32−, SO32−, SO42− and polythionates [16]. SO42− applied as fertilizer or obtained through the processes described above is the form of S that plants assimilate through their roots [40].
At best growth conditions, a plant’s sulfur requirement ranges from 2 to 10 μmol g−1 plant fresh weight day−1 [1]. As the flow of S is a dynamic process where the ecosystem receives S from the atmosphere, precipitation, subsoil water, and fertilizers, and loses S through the process of volatilization of S by soil and plants and by leaching, it is difficult to estimate the actual amount of S that a plant surface absorbs, assimilates, leaches, and volatilizes. As an exercise, let us suppose a single sampling point for a field of maize, for example before the harvest. In one hectare, there may be 78,000 kg of fresh weight ha−1, which would be equivalent to 25 kg of sulfur contained in the plants. However, the 25 kg ha−1 accumulated in the plant tissues at that specific sampling time does not include the S volatilized by the plant itself, or that leached, assimilated, or volatilized in the soil and by the microorganisms.
The point to highlight with the data of the previous paragraph is that the S of the soil is in constant exchange and extraction by the crops, atmosphere, and soil water. Therefore, a continuous supply of S is required, which is recommended to be applied in the form of S0 (40–60 kg ha−1) every one or two years, to maintain the edaphic store.

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Subjects: Agronomy
Contributors MDPI registered users' name will be linked to their SciProfiles pages. To register with us, please refer to https://encyclopedia.pub/register :
Laura Olivia Fuentes-Lara
,
Julia Medrano-Macías
,
Fabián Pérez-Labrada
,
Erika Nohemí Rivas-Martínez
,
Ema Laura García-Enciso
,
Susana González-Morales
,
Antonio Juárez-Maldonado
,
Froylán Rincón-Sánchez
,
Adalberto Benavides-Mendoza
View Times: 141
Revisions: 2 times (View History)
Update Date: 20 Nov 2023
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