Solute accumulation of chestnut oak and dogwood leaves in response to throughfall manipulation of an upland oak forest

To determine the biochemical basis of osmotic adjustment, seasonal and treatment differences in foliar water-soluble organic solutes and inorganic ions were investigated for two hardwood species that exhibited osmotic adjustment in a Throughfall Displacement Experiment at the Walker Branch Watershed near Oak Ridge, Tennessee. Leaf samples of overstory and understory chestnut oak (Quercus prinus L.) and understory dogwood (Cornus florida L.) were collected during the 1994 (wet) and 1995 (dry) growing seasons from each of three treatments: dry (-33% throughfall), ambient (control) and wet (+33% throughfall). Foliar soluble carbohydrates and organic acids were measured by gas chromatography-mass spectrometry. During May 1994, when the demand for sucrose was greatest, dogwood accumulated small amounts of glucose, quinic acid and Mg²⁺, offsetting a decline in nitrate concentration. As the mild drought continued and tree growth slowed, there was a significant accumulation of sucrose in dogwood in the dry treatment in June, and a trend toward increased K⁺. In overstory chestnut oak in the dry treatment over the same period, there were significant accumulations of fructose, glucose and K⁺, and a trend toward increased quinic acid accumulation. Sucrose did not become a key osmotically active compound in chestnut oak until August 1994. In 1995, with the exception of understory chestnut oak, there was no accumulation of K⁺ in either species. During the severe drought of 1995, monosaccharides, particularly glucose and fructose, accounted for most of the osmotic adjustment in both species. Among solutes, glucose constituted the largest accumulation in dogwood in the dry treatment in August 1995, followed by fructose and sucrose. There was only a moderate increase in solutes in chestnut oak trees in 1995, with fructose and glucose constituting over 50% of the predicted solute accumulation in July. Both species accumulated a wider array of solutes during the dry year than during the wet year, but treatment differences in solute accumulation in chestnut oak were partially limited by drought. The greater dehydration tolerance of chestnut oak than dogwood was evident in the higher baseline concentrations of organic solutes and the greater array of solutes accumulated in response to drought.