The combined effects of fertilization and relative water limitation on tissue water relations, hydraulic parameters and shallow root distribution in loblolly pine (Pinus taeda L.)

Abstract

One goal of this research was to characterize shoot tissue-level responses in loblolly pine to soil moisture limitation in combination with fertilization as well as to more severe soil moisture limitation. We found that neither fertilization alone, nor fertilization in combination with soil moisture limitation resulted in changes to shoot tissue water relations parameters classically characterized in drought response studies. More severe water limitation was necessary to elicit responses, and those responses had not been fully described previously. The more severe water limitation resulted in increased capacitance beyond turgor loss, increased relative water content at turgor loss, a more negative turgor loss point, an increased bulk modulus of elasticity, more negative osmotic potential at 100% relative water content, and an increased apoplastic water fraction. As there were indications of reduced water use and moisture stress in the absence of shoot level responses under less severe drought, such parameters are insufficient alone to characterize moisture stress in fertilized and in less severely water limited loblolly trees. Additionally, we sought a morphological or physiological explanation for the reduced transpiration and increased water use efficiency reported for fertilized trees in the Virginia Piedmont. Our characterizations of the responses of root distribution and hydraulics to limited soil moisture here complement existing research, which demonstrated changes to root distribution and hydraulics in response to fertilization. The responses we discovered in fertilized trees that accompanied reduced transpiration and increased water use efficiency that differed from responses to reduced soil moisture alone were primarily large decreases to shallow root presence. We found this to be readily quantified using measures of root length density. Decreases to whole-tree hydraulic conductivity were also shown to occur with fertilization and were shown not to occur in shoot tissue, suggesting limitation via rhizosphere or root xylem conductance. Our results support the supposition that fertilization narrows hydraulic safety margins and potentially predisposes loblolly trees to moisture stress, particularly prolonged, severe water limitation following fertilization. Finally, we tested the validity of throughfall exclusion for simulating reduced rainfall using a greenhouse 'split-pot' study, which applied spatially fixed heterogeneous soil moisture to young, well-watered loblolly pines. The 'split-pot' experiments demonstrated that spatially fixed soil moisture heterogeneity does not confound drought effects; needle area specific transpiration was not decreased, nor was water use efficiency increased. This supports the validity of inferences taken from drought simulation experiments with loblolly pine where throughfall exclusion troughs reduce soil moisture content in a consistent, spatially heterogeneous manner.