Metabolic responses of Norway spruce (Picea abies) trees to long-term forest management practices and acute (NH₄)₂SO₄ fertilization: transport of soluble non-protein nitrogen compounds in xylem and phloem

Abstract

From laboratory experiments with seedlings and young trees of Norway spruce (Picea abies L. Karst.), a cycling pool of soluble non-protein N compounds is thought to be indicative of the N-nutritional status of trees. In order to test whether this assumption can be transferred to mature trees grown in the field, xylem sap and phloem exudate were collected from spruce trees in two remote forest stands: (1) a N-limited stand (Villingen site), and (2) a stand where trees are sufficiently supplied with N from the soil (Schluchsee site). Trees at these sites were c. 80–100 (Villingen site) and c. 40–60 (Schluchsee site) yr old. In addition to untreated control areas, one entire watershed area at both sites was subjected to (NH₂)₂SO₄ fertilization to the soil.

In the xylem sap of the spruce roots at both sites Gln, Asp and Arg were the dominant total soluble non-protein nitrogen (TSNN) compounds. In the xylem sap of the trunk and the twigs Arg was virtually absent and Gln plus Asp dominated TSNN. On average, TSNN in the xylem sap of trees at the Schluchsee site was 1·5–2-fold higher than those at Villengen. Highest TSNN contents in the xylem sap were found during growth and development of current year tissues, while the lowest TSNN contents were found in summer. At the Villingen site (NH₄)₂SO₄ fertilization caused an increase in the Gln content in the xylem sap of all tree sections analysed as well as an increase in the Arg content in the xylem sap of the roots. At the Schluchsee site only a small increase in TSNN contents of the xylem was observed, mostly in the xylem sap of the roots. In phloem exudates TSNN contents were much higher in trees on the Schluchsee than on the Villingen site. The seasonal pattern of TSNN in phloem exudates was similar to the seasonal pattern found in the xylem sap. During spring and early summer Gln was the predominant TSNN compound in phloem exudates, but during late summer and autumn Arg became predominant. At the Villingen site (NH₄)₂SO₄ fertilization caused a significant increase in TSNN contents in phloem exudates of twigs and roots, but at Schluchsee an increase in TSNN was found only in phloem exudates of the roots. At both field sites Arg that was not transported to the shoot by xylem transport, was allocated from the leaves to the roots by phloem transport and was cycled within the root system by both xylem and phloem transport. From these results it is calculated that shoot-to-root signalling by long-distance transport of amino compounds can also contribute to the regulation of N-nutrition of mature spruce trees. Apparently, the internal cycling of individual N compounds within spruce trees differs considerably.