References on sulfur use are found in the Bible and in Greek and Roman literature. As early as more than 2000 years ago, Romans discovered the beneficial effects of sulfur as a potent antidote against plant pathogens and refer the application of elemental sulfur in the vineyards (Rausch 2007). To date, the reports on SO42- nutrition or supply or symptoms of sulfur deficiency in grapevine are scarce.
The effect of sulfur on plant growth, productivity and product quality mostly relates sulfur nutrition in interaction with nitrogen (Brunold 1976, Byers et al. 1987, Schnug 1997). In grapevine xylem sap, sulfate, chloride and phosphate increased by N fertilizer treatment, while nitrate was the major anion followed by malate (Peuke 2000). The protective effect of elemental S against pests and diseases has been mostly reported after foliar application (Bloem et al. 2007). In the decade of 90's field trials showed that S fertilization could substitute for fungicide application in crop yield protection from pests attack. Then the concept of Sulfur Induced Resistance (SIR) was introduced to describe the natural resistance of plants against fungal pathogens through the stimulation of sulfur metabolic processes by fertilizer application strategies (Haneklaus et al. 2007), although S-containing metabolites responsible for SIR have not been fully elucidated (Bloem et al. 2007).
In grapevine organic farming, incorporation into the soil of fine-granule sulfur and Thiobacillus microorganisms, restricted powdery mildew infected berries and leaves by more than 80%, in comparison with 90% after treatment with a conventional fungicide used as control (Bourbos et al. 2000). In fact, after the introduction of powdery mildew (Erysiphe necator, previously Uncinula necator) in Europe, in the XIX century, it was noticed that sulfur applied to vine leaves and berries significantly protected from powdery mildew infection. Tra ditionally, treatments against this disease used copper sulfate as Bordeaux mixture, a high S content fungicide, or S0 applied as a dust or a wetable powder (Williams and Cooper 2004). Aqueous solution of lime sulfur was also used as a spray and when applied to dormant grapevines it delayed the epidemic development of powdery mildew (Gadoury et al. 1994). CuSO4 alone, especially at low concentrations, elicited a substantial production of phytoalexins in grapevine leaves. Evidence is also provided that chitosan (CHN), a deacetylated derivative of chitin, which significantly reduces the infection of grapevine leaves by Botrytis cinerea and Plasmopara viticola, in combination with CuSO4 confers protection against both (Aziz et al. 2006). S0 has been used with caution due to eventual inhibition of the microbial activity during the grape must fermentation and residues remaining in the wine (Schutz and Kunkee 1977).
However, residual sulfur levels decrease rapidly following the last dusting (Thomas et al. 1993) and late S0 application does not interfere with alcoholic fermentation neither affects wine analytical parameters (Jolivet 1993). Despite the advantages of elemental sulfur due to its target activity against powdery mildew, low cost and organic production, vineyard fungicides are increasingly substituted for chemicals without any sulfur; besides, last generation fertilizers contain low S contamination. Nevertheless, many compounds responsible for typical aromas of Vitis vinifera wines are sulfur containing substances (dePinho et al. 1997).
The availability of sulfur in the vineyards, grapevine sulfate nutrition and sulfur as biotic stress antagonist are important topics for separate review articles. In the present review we report advances in the last few years on the expression and regulation of Vitis genes encoding for sulfur transporters and sulphur assimilating enzymes.
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