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Revisiting Sulphur—The Once Neglected Nutrient: It's Roles in Plant Growth, Metabolism, Stress Tolerance and Crop Production

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Revisiting Sulphur—The Once Neglected Nutrient: It's Roles in Plant Growth, Metabolism, Stress Tolerance and Crop Production agriculture Review Revisiting Sulphur—The Once Neglected Nutrient: It’s Roles in Plant Growth, Metabolism, Stress Tolerance and Crop Production Tinashe Zenda 1,2,3,4 , Songtao Liu 1,2,3, Anyi Dong 1,2,3 and Huijun Duan 1,2,3,* 1 State Key Laboratory of North China Crop Improvement and Regulation, Hebei Agricultural University, Baoding 071001, China; [email protected] (T.Z.); [email protected] (S.L.); [email protected] (A.D.) 2 North China Key Laboratory for Crop Germplasm Resources of the Education Ministry, Hebei Agricultural University, Baoding 071001, China 3 Department of Crop Genetics and Breeding, College of Agronomy, Hebei Agricultural University, Baoding 071001, China 4 Department of Crop Science, Faculty of Agriculture and Environmental Sciences, Bindura University of Science Education, Bindura P. Bag 1020, Zimbabwe * Correspondence: [email protected]; Tel.: +86-139-3127-9716 Abstract: Sulphur plays crucial roles in plant growth and development, with its functions ranging from being a structural constituent of macro-biomolecules to modulating several physiological processes and tolerance to abiotic stresses. In spite of these numerous sulphur roles being well acknowledged, agriculture has paid scant regard for sulphur nutrition, until only recently. Serious problems related to soil sulphur deficiencies have emerged and the intensification of food, fiber, and animal production is escalating to feed the ever-increasing human population. In the wake of huge Citation: Zenda, T.; Liu, S.; Dong, A.; demand for high quality cereal and vegetable diets, sulphur can play a key role in augmenting the Duan, H. Revisiting Sulphur—The production, productivity, and quality of crops. Additionally, in light of the emerging problems of soil Once Neglected Nutrient: It’s Roles in fertility exhaustion and climate change-exacerbated environmental stresses, sulphur assumes special Plant Growth, Metabolism, Stress importance in crop production, particularly under intensively cropped areas. Here, citing several Tolerance and Crop Production. Agriculture 2021, 11, 626. https:// relevant examples, we highlight, in addition to its plant biological and metabolism functions, how doi.org/10.3390/agriculture11070626 sulphur can significantly enhance crop productivity and quality, as well as acclimation to abiotic stresses. By this appraisal, we also aim to stimulate readers interests in crop sulphur research by Academic Editors: Pavel Krasilnikov, providing priorities for future pursuance, including bettering our understanding of the molecular Miguel A. Taboada and Amanullah processes and dynamics of sulphur availability and utilization in plants, dissecting the role of soil rhizospherical microbes in plant sulphur transformations, enhancing plant phenotyping and Received: 28 May 2021 diagnosis for nutrient deficiencies, and matching site-specific crop sulphur demands with fertilizer Accepted: 29 June 2021 amendments in order to reduce nutrient use inefficiencies in both crop and livestock production Published: 3 July 2021 systems. This will facilitate the proper utilization of sulphur in crop production and eventually enhance sustainable and environmentally friend food production. Publisher’s Note: MDPI stays neutral with regard to jurisdictional claims in Keywords: sulphur; crop productivity; crop quality; sulphate transporters; secondary sulphur published maps and institutional affil- compounds; abiotic stress tolerance; crop production iations. 1. Introduction Copyright: © 2021 by the authors. Licensee MDPI, Basel, Switzerland. Realizing the mounting global demand for adequate and highly nutritious food as This article is an open access article well as fiber for the rising human population, there is a need to increase global crop produc- distributed under the terms and tion [1]. One way for improving crop production and productivity is through modifying conditions of the Creative Commons soil nutrient supply, among other strategies [2]. Sulphur, (S) macronutrient, essential for Attribution (CC BY) license (https:// plant growth and development [3–7], has also been acknowledged for improving crop creativecommons.org/licenses/by/ productivity [8–13], quality [14–16], and plants’ abiotic stress responses [17–22]. Therefore, 4.0/). access to an adequate supply of S for plants throughout their development is necessary for Agriculture 2021, 11, 626. https://doi.org/10.3390/agriculture11070626 https://www.mdpi.com/journal/agriculture Agriculture 2021, 11, 626 2 of 24 optimum crop performance [23]. Despite all this, however, S macronutrient has received little attention for many years, until only recently [17,24]. This is largely because, previ- ously, it was thought fertilizers and atmospheric deposition adequately supplied the soil with enough of the nutrient [9,25]. Consequently, little crop research on S nutrient has been conducted and very scarce information has been reported [9,17]. However, remarkable revelations indicate that most arable soil around the world has been deficient in S for over three decades, but the negative effects on global crop production have only recently become evident [4,12,26]. In recent years, diminished atmospheric S deposition as a result of stricter emission regulations, decreased use of S based fungicides, use of high analysis low S synthetic fertilizers, low S returns from farm animal derived manure, cultivation of high nutrient-demanding and high yielding crop cultivars, and intensification of agriculture have all led to widespread S shortage across the globe [17]. Subsequently, due to the importance of S nutrient in plant functions (such as in sugar production, carbon dioxide assimilation, nitrogen (N) fixation and protein formation), S nutrient is increasingly becoming more important [13]. Notably, scientific research indicates that the crop farmers will have to start applying S nutrient through fertilization to achieve optimal production from ever-improving, high performance genetics [24]. Thus, the re-emerging interest in S has recently spurred its global demand. In terms of market, the need for S fertilizers is increasing at a considerable rate [27], with the global S fertilizer market accounting for US $4.52 billion in 2017 and expected to reach US $7.03 billion by 2026 [28]. This is largely driven by an upsurge in demand in the agriculture sector for crops prone to S deficiency and need for higher productivity. In addition, the degradation of soil quality and increase in S deficiency in the soil, reduction in S emission influencing the requirement for added S, surge in use of sulphur in fungicides and insecticides are the drivers to the growth of the S market [28]. By type, elemental S segment is expected to be the fastest growing during the forecast period. Particularly, the use of elemental S is increasing due to its ability to reduce soil pH and to reclaim sodic soils. By crop type, the oilseeds and pulses (especially canola and soybean (Glycine max L.) segment will dominate the high S requirements, largely owing to the fact that S enhances oil synthesis and protein production in pulses [29]. Further projections show that the Asia Pacific region will be the fastest-growing market during the forecast period 2017–2026, being driven by the increase in agricultural practices and the requirement for high-quality agricultural produce [30], with the Chinese fertilizer demand playing a major role [31]. In crop production, S is attracting the most attention in soil science and plant nutrition due to its good nutritive potentiality to crops, its potential protective role to abiotic stresses, and its relative immobility in the soil–plants system [32,33]. It has been observed that, without adequate S, crops cannot reach their full potential in terms of yield, quality or protein content, nor can they make efficient use of applied N, caused by the S requirement for protein and enzyme synthesis [4,10,17,34]. In the wake of huge demand for high quality cereal and vegetable diets, S can play a key role in augmenting the production, productivity, and quality of crops [35,36]. Moreover, in light of emerging problems of soil fertility exhaustion, S assumes special importance for crop production, particularly under intensively cropped areas. In this review, therefore, we discuss the role of S nutrient in crop production, particularly focusing on plant growth, metabolism and acclimation to abiotic stresses, as well as improving crop productivity. We then conclude by highlighting the future prospects of S research in crops. 2. Biological Functions of Sulphur (S) in Plants Sulphur is an essential macronutrient, vital for plant growth and functioning; ranked fourth after nitrogen, phosphorus (P) and potassium (K); with total S content in plant tissues ranging from 0.3% to 7.6% [3,36]. Precisely, S is of great importance for the primary structure of proteins and functioning of enzymes, by being a constituent of the amino acids cysteine (Cys) and methionine (Met) [21,37–40]. The structural compositions of these essential S containing amino acids are shown in Figure1. Agriculture 2021, 11, x FOR PEER REVIEW 3 of 25 Sulphur is an essential macronutrient, vital for plant growth and functioning; ranked fourth after nitrogen, phosphorus (P) and potassium (K); with total S content in plant tissues ranging from 0.3% to 7.6% [3,36]. Precisely, S is of great importance for the primary structure of proteins and functioning of enzymes, by being a constituent of the Agriculture 2021, 11, 626 3 of 24 amino acids cysteine (Cys) and methionine
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