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The effect of iron (Fe) and sulphur (S) deprivation on sulphate uptake and assimilation pathways was investigated in durum wheat by analysing the expression of genes coding for major transporters and enzymes involved in sulphate assimilation and reduction: high-affinity sulphate transporters (TdSultr1.1 and TdSultr1.3), ATP sulphurylase (TdATPSul1 and TdATPSul2), APS reductase (TdAPR), sulphite reductase (TdSiR), O-acetylserine(thiol)lyase (TdOASTL1 and TdOASTL2), and serine acetyltransferase (TdSAT1 and TdSAT2). Further experiments were carried out to detect changes in the activities of these enzymes, together with the evaluation of growth parameters (fresh biomass accumulation, leaf green values, and total S, thiol, and Fe concentrations). Fe shortage in wheat plants under adequate S nutrition resulted in an S deficiency-like response. Most of the genes of the S assimilatory pathway induced by S deprivation (TdATPSul1, TdAPR, TdSir, TdSAT1, and TdSAT2) were also significantly up-regulated after the imposition of the Fe limitation under S-sufficient conditions. However, the differential expression of genes encoding the two high-affinity transporters (TdSultr1.1 and TdSultr1.3) indicates that the mechanisms of sulphate uptake regulation under Fe and S deficiency are different in wheat. Moreover, it was observed that the mRNA level of genes encoding ATPS, APR, and OASTL and the corresponding enzyme activities were often uncoupled in response to Fe and S availability, indicating that most probably their regulation involves a complex interplay of transcriptional, translational, and/or post-translational mechanisms induced by S and/or Fe deficiency.

Characterization of the relationship between sulfur and iron in both Strategy I and Strategy II plants, has proven that low sulfur availability often limits plant capability to cope with iron shortage. Here it was investigated whether the adaptation to iron deficiency in tomato (Solanum lycopersicum L.) plants was associated with an increased root sulfate uptake and translocation capacity, and modified dynamics of total sulfur and thiols accumulation between roots and shoots. Most of the tomato sulfate transporter genes belonging to Groups 1, 2, and 4 were significantly upregulated in iron-deficient roots, as it commonly occurs under S-deficient conditions. The upregulation of the two high affinity sulfate transporter genes, SlST1.1 and SlST1.2, by iron deprivation clearly suggests an increased root capability to take up sulfate. Furthermore, the upregulation of the two low affinity sulfate transporter genes SlST2.1 and SlST4.1 in iron-deficient roots, accompanied by a substantial accumulation of total sulfur and thiols in shoots of iron-starved plants, likely supports an increased root-to-shoot translocation of sulfate. Results suggest that tomato plants exposed to iron-deficiency are able to change sulfur metabolic balance mimicking sulfur starvation responses to meet the increased demand for methionine and its derivatives, allowing them to cope with this stress.

It is well known that S interacts with some macronutrients, such as N, P, and K, as well as with some micronutrients, such as Fe, Mo, Cu, Zn, and B. From our current understanding, such interactions could be related to the fact that: (i) S shares similar chemical properties with other elements (e.g., Mo and Se) determining competition for the acquisition/transport process (SULTR transporter family proteins); (ii) S-requiring metabolic processes need the presence of other nutrients or regulate plant responses to other nutritional deficiencies (S-containing metabolites are the precursor for the synthesis of ethylene and phytosiderophores); (iii) S directly interacts with other elements (e.g., Fe) by forming complexes and chemical bonds, such as Fe-S clusters; and (iv) S is a constituent of organic molecules, which play crucial roles in plants (glutathione, transporters, etc.). This review summarizes the current state of knowledge of the interplay between Fe and S in plants. It has been demonstrated that plant capability to take up and accumulate Fe strongly depends on S availability in the growth medium in both monocots and dicot plants. Moreover, providing S above the average nutritional need enhances the Fe content in wheat grains, this beneficial effect being particularly pronounced under severe Fe limitation. On the other hand, Fe shortage induces a significant increase in the demand for S, resulting in enhanced S uptake and assimilation rate, similar to what happens under S deficiency. The critical evaluation of the recent studies on the modulation of Fe/S interaction by integrating old and new insights gained on this topic will help to identify the main knowledge gaps. Indeed, it remains a challenge to determine how the interplay between S and Fe is regulated and how plants are able to sense environmental nutrient fluctuations and then to adapt their uptake, translocation, assimilation, and signaling. A better knowledge of the mechanisms of Fe/S interaction might considerably help in improving crop performance within a context of limited nutrient resources and a more sustainable agriculture.

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