Effects of humic substances on carbohydrate metabolism of maize leaves
Maize plants were grown in nutrient solution with or without 100 mg organic carbon L−1 of humic substances (HS) (MW > 12 000) extracted from a cambisol. To evaluate the effect of HS on leaf carbohydrate metabolism, after 14 d of growth apical and basal segments from the third leaf were collected and analyzed for sugar content and for the activities of enzymes involved in the synthesis and breakdown of starch and sucrose. Leaf starch content decreased in treated plants, whereas the level of soluble sugars increased. The decrease of starch was accompanied by an enhanced activity of amylase, whereas the activity of ADPglucose pyrophosphorylase was not affected. Activities of invertases and sucrose synthase were stimulated by HS treatment in apical tissue, and inhibited in basal one. Activities of enzymes involved in sucrose biosynthesis (sucrose phosphate synthase, cytoplasmic fructose-1,6- bisphosphatase, UDPglucose pyrophosphorylase) were stimulated or inhibited depending on the enzyme and on the tissue examined. Our results show that HS may affect both the level and percent distribution of sugars of maize leaves; these effects seem to be mediated by changes in the activities of enzymes involved in carbohydrate metabolism. Key words: Humic substances, enzymes, carbohydrate metabolism, maize