Determinants of legacy effects in pine trees - implications from an irrigation-stop experiment.

Tree responses to altered water availability range from immediate (e.g. stomatal regulation) to delayed (e.g. crown size adjustment). The interplay of the different response times and processes, and their effects on long-term whole-tree performance, however, is hardly understood. Here we investigated legacy effects on structures and functions of mature Scots pine in a dry inner-Alpine Swiss valley after stopping an 11-yr lasting irrigation treatment. Measured ecophysiological time series were analysed and interpreted with a system-analytic tree model. We found that the irrigation stop led to a cascade of downregulations of physiological and morphological processes with different response times. Biophysical processes responded within days, whereas needle and shoot lengths, crown transparency, and radial stem growth reached control levels after up to 4 yr only. Modelling suggested that organ and carbon reserve turnover rates play a key role for a tree's responsiveness to environmental changes. Needle turnover rate was found to be most important to accurately model stem growth dynamics. We conclude that leaf area and its adjustment time to new conditions is the main determinant for radial stem growth of pine trees as the transpiring area needs to be supported by a proportional amount of sapwood, despite the growth-inhibiting environmental conditions.

Plasticity in gas-exchange physiology of mature Scots pine and European larch drive short- and long-term adjustments to changes in water availability.

Adjustment mechanisms of trees to changes in soil-water availability over long periods are poorly understood, but crucial to improve estimates of forest development in a changing climate. We compared mature trees of Scots pine (Pinus sylvestris) and European larch (Larix decidua) growing along water-permeable channels (irrigated) and under natural conditions (control) at three sites in inner-Alpine dry valleys. At two sites, the irrigation had been stopped in the 1980s. We combined measurements of basal area increment (BAI), tree height and gas-exchange physiology (Δ13 C) for the period 1970-2009. At one site, the Δ13 C of irrigated pine trees was higher than that of the control in all years, while at the other sites, it differed in pine and larch only in years with dry climatic conditions. During the first decade after the sudden change in water availability, the BAI and Δ13 C of originally irrigated pine and larch trees decreased instantly, but subsequently reached higher levels than those of the control by 2009 (15 years afterwards). We found a high plasticity in the gas-exchange physiology of pine and larch and site-specific responses to changes in water availability. Our study highlights the ability of trees to adjust to new conditions, thus showing high resilience.

Trait-specific responses of Scots pine to irrigation on a short vs long time scale.

In xeric environments, an increase in drought is related to reduced forest productivity and to enhanced mortality. However, predictions of future forest development remain difficult as the mechanisms underlying the responses of mature trees to long-term variations in water availability are not well understood. Here, we aimed to compare the adjustments in radial growth and morphological needle and shoot traits of mature Scots pine (Pinus sylvestris L.) growing along open water channels with those of control trees growing under naturally dry conditions at three sites in Valais, an inner-Alpine dry valley of Switzerland. The trees growing along two channels had been irrigated since germination (>70 years), whereas those along another previously drained channel had been irrigated only from 2010 to 2012, when the channel was re-established, and could thus be used to quantify the short-term effects of re-irrigation. Linear mixed models revealed that needle and shoot lengths as well as early- and late-wood basal area increments (BAIs) were most responsive to short-term and long-term irrigation. However, the magnitude of the response to the short-term irrigation exceeded that of the long-term irrigation. An extreme drought during the first half of 2011 led to an immediate decrease in the needle length, needle width, and early- and late-wood BAIs of the control trees, whereas the shoot length and needle numbers of control trees reacted with a 1-year delay to the extreme drought, as the shoots were responding to water availability of previous year's summer. Such negative responses to dry climatic conditions were even found in irrigated trees at one of our sites, which might be linked to tree growth becoming more sensitive to drought with increasing tree height and leaf area. In order to improve predictions of future forest development, long-term studies are necessary that consider lagged responses and adjustment processes of trees to changes in water availability.