Abstract
Tree saplings are exposed to a competitive growth environment in which resources are limited and the ability to adapt determines general vitality and specific growth performance. In this study we analyzed the aboveground spatial neighborhood of oak [Quercus petraea (Matt.) Liebl.] and beech (Fagus sylvatica L.) saplings growing in Germany, by using hemispherical photography and terrestrial laser scanning as proxy for the competitive pressure saplings were exposed to. The hemispherical images were used to analyze the light availability and the three-dimensional (3D) point clouds from the laser scanning were used to assess the space and forest structure around the saplings. The aim was to increase the precision with which the biomass allocation, growth, and morphology of the saplings could be predicted by including more detailed information of their environment. The predictive strength of the models was especially increased through direct neighborhood variables (e.g., relative space filling), next to the light availability being the most important predictor variable. The biomass allocation patterns within the more light demanding oak were strongly driven by the space availability around the saplings. Diameter and height growth variables of both species reacted significantly to changes in light availability, and partly also to the neighborhood variables. The leaf morphology [as leaf-area ratio (LAR)] was also driven by light availability and decreased with increasing light availability. However, the branch morphology (as mean branch weight) could not be explained for oak and the model outcome for beech was hard to interpret. The results could show that individuals of the same species perform differently under constant light conditions but differing neighborhoods. Assessing the neighborhood of trees with highly precise measurement devices, like terrestrial laser scanners, proved to be useful. However, the primary response to a dense neighborhood seemed to be coping with a reduction of the lateral light availability aboveground, rather than responding to an increase of competition belowground. The results suggest continuing efforts to increase the precision with which plant environments can be described through innovative and efficient methods, like terrestrial laser scanning.
Highlights
Every plant competes with other plants or organisms for limited resources
Effects of belowground competition on sapling performance induced by mature trees have been shown by exclusionary experimental setups, e.g., root-trenching experiments (Leuschner et al, 2001; Ammer, 2002; Petritan et al, 2011), even though still comparably little is known about belowground competition
All other models had more than one predictor variable, with their relative importance ranging from 0.03 up to 0.52, respectively
Summary
Every plant competes with other plants or organisms for limited resources. The survival and general plant performance is primarily determined by the amount of resources the individual plant can capture. For saplings growing in the understory of (temperate) forest ecosystems the amount of light is considered to be among the most important resources (Lambers et al, 2008). The availability of light is known to influence different species in various ways, especially when considering light demanding or shade tolerant species (Poorter, 1999; Lödige et al, 2014; Annighöfer et al, 2017), resulting in species-specific traits as reaction to varying levels of light (Petritan et al, 2009; Schall et al, 2012). Plants in general compete for other resources above- and belowground, including water, nutrients and growing space (Casper and Jackson, 1997; Leuschner and Ellenberg, 2017). Effects of belowground competition on sapling performance induced by mature trees have been shown by exclusionary experimental setups, e.g., root-trenching experiments (Leuschner et al, 2001; Ammer, 2002; Petritan et al, 2011), even though still comparably little is known about belowground competition
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