Research Summary• We tested whether the degree of shade-induced plasticity in petiole length and leaf area is related to the mean trait value expressed under high-light conditions, and to what extent trait values expressed under high-light and shaded conditions affect plant performance.• Thirty-four Trifolium repens genotypes were used with a wide range of petiole lengths and leaf areas. Plants were subjected to a high-light environment and two shading regimes: homogeneous shading and a vertical light gradient.• Absolute petiole elongation in response to both shading treatments and absolute leaf area expansion in response to homogeneous shading were independent of the trait values expressed in high light. Consequently, relative plasticity was higher for genotypes with lower high-light trait values. Plasticity was associated with enhanced plant performance in a vertical light gradient but not in homogeneously shaded conditions. We also found costs associated with the ability to express plasticity.• Our results suggest that selection can act separately on trait values expressed under high-light conditions and on the degree of plasticity.

We studied the effects of genotypic and plastic variation in vertical and horizontal spacer lengths on plant performance in a stoloniferous herb subjected to opposing selection regimes. We hypothesized that longer vertical structures are beneficial if plants are subjected to competition, but they should negatively affect plant performance if plants are exposed to aboveground disturbance. To test these hypotheses we subjected 34 genotypes of Trifolium repens to competition and disturbance treatments. Competition was imposed by a grass canopy consisting of Lolium perenne, and disturbance was simulated by regularly clipping the target plants and all the surrounding vegetation at 1 cm above soil level. Conform to our hypothesis, genotypes with longer vertical structures (petioles) produced fewer ramets than genotypes with shorter petioles in the disturbance treatment. However, genotypes with longer petioles did not perform better under competition than genotypes with shorter petioles. Genotypes with highly plastic vertical structures tended to produce more shoot mass under competition, and they produced fewer ramets if subjected to disturbance. Unexpectedly, horizontal structures (stolon internodes) expanded in response to competition which, furthermore, was associated with enhanced plant performance. However, producing longer internodes is inherently associated with costs in terms of increased resource allocation to the longer structures, but not to benefits in terms of increased resource capture. Positive correlations among the length and plasticity of vertical and horizontal structures may explain the apparent positive effect of producing longer internodes on plant performance. Our data thus support the notion that trait correlations may weaken selective forces acting on a focal trait in a specific environment if opposing selection pressures act on genetically correlated traits.

In stoloniferous species, the length of petioles is of pivotal importance because it determines the position of leaf blades within the canopy. From a mechanistic perspective, two developmental processes, cell division and cell elongation, are responsible for the length of a given petiole. This study aimed at quantifying the relative contributions of cell division and cell elongation to genotypic and plastic variation in petiole length of the stoloniferous herb Trifolium repens. Thirty-four genotypes of T. repens were grown under high light conditions and simulated canopy shade. Cells were counted and their lengths measured on epidermal prints from fully grown petioles of leaves that had been initiated in the experimental light conditions. Cell number was the main trait explaining petiole length differences among genotypes grown under high light, while both cell number and length changed in response to shading. Our study revealed a strong negative correlation between shade-induced changes in cell number and cell length: genotypes that responded to shading by increasing cell numbers hardly changed in cell length, and vice versa. Our results suggest that genotypic and phenotypic variation in petiole length results from a complex interplay between the developmental processes of cell elongation and cell division.

Increased cell number and cell length both contribute to shade induced elongation of petioles which enables stoloniferous plants to place their leaf lamina higher up in the canopy. Although petiole elongation is assumed to be beneficial, it may also imply costs in terms of decreased biomechanical stability. We test the hypothesis that shade induced elongation changes the biomechanical properties of petioles and that the underlying mechanisms, cell division and cell elongation, differentially affect biomechanical properties. This was done by subjecting 14 genotypes differing in the relative contribution of cell size and cell number to shade induced elongation responses to high light conditions and to simulated canopy shade. Developmental traits (cell size and cell number), morphological traits characterizing the petioles, as well as biomechanical characteristics were measured. Our results show that, comparable to stems of non-clonal plants, the rigidity of a petiole's tissue (the Young's modulus) increases, leading to increased flexural stiffness of petioles subjected to shading. Increased flexural stiffness proved to be associated with increased performance under shaded conditions. Our results also indicate that cell number affected the material properties and the flexural stiffness of petioles. However, the degree and pattern of the effects differed between light environments. Shade induced increase in cell number translated into shade induced increase of Young's modulus and flexural stiffness. Genotypes producing relatively larger cells under shaded conditions experienced a decrease in tissue rigidity. In concert our results indicate that the pattern of selection on flexural stiffness, and thereby also on shade induced changes of cell number and cell size differs among light environments.

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