Excess of exogenous nitrates inhibits formation of abnormal wood in the Karelian birch

Abstract

The effect of exogenous nitrate on the sucrose-metabolizing enzyme activities—sucrose synthase (SS) and apoplastic invertase (ApInv)—in the xylem and phloem of the silver (Betula pendula Roth var. pendula) and Karelian (B. pendula var. carelica) birches (the latter is well known for its abnormal, patterned wood) has been studied. A stable correlation between the enzyme activities and deviations in the growth and development of stem vascular tissues during cambial growth has been demonstrated. Formation of the birch wood with a common structure is associated with high SS activity. In this case, the reaction yields UDP glucose, which is utilized mainly for synthesis of the cell wall components of vessels and fiber tracheids. As for the Karelian birch, the SS activity in the xylem formation zone is decreased, which complies with a higher sucrose level in the tissue. The excess sucrose is released into the apoplast to be cleaved by ApInv. The resulting hexoses induce storage metabolism, thereby increasing the amount of storage substances and the share of storage parenchyma cells in the xylem. As a result, the Karelian birch wood acquires large inclusions in the parenchyma, which render a characteristic pattern. A change in the ratio of SS to ApInv activities underlies a great variety in the degree of wood patterning observed in Karelian birch trees. In the common silver birch, the nitrate application increases the sucrose utilization via SS pathway, which results in an increase in wood growth. In the Karelian birch xylem, nitrates lead to a decrease in both the SS (a decrease in wood growth) and ApInv (a decrease in the amount of parenchyma, i.e., normalization of the wood structure). The sucrose metabolizing in the xylem decreases on the background of an increase in its utilization in the phloem, where both enzyme activities elevate. It is assumed that the fact that the Karelian birch distribution range is limited by rich soils can be determined by a shift from intensive apoplastic sucrose utilization zone towards the phloem caused by high doses of nitrogen nutrition.