The role of biomass allocation in the growth response of plants to different levels of light, CO 2 , nutrients and water: a quantitative review

The allocation of biomass to different plant organs depends on species, ontogeny and on the environment experienced by the plant. In this paper we first discuss some methodological tools to describe and analyse the allocation of biomass. Rather than the use of shoot:root ratios, we plead strongly for a subdivision of biomass into at least three compartments: leaves, stems and roots. Attention is drawn to some of the disadvantages of allometry as a tool to correct for size differences between plants. Second, we tested the extent to which biomass allocation of plants follows the model of a ‘functional equilibrium’. According to this model, plants respond to a decrease in above-ground resources with increased allocation to shoots (leaves), whereas they respond to a decrease in below-ground resources with increased allocation to roots. We carried out a meta-analysis of the literature, analysing the effect of various environmental variables on the fraction of total plant biomass allocated to leaves (leaf mass fraction), stem (stem mass fraction) and roots (root mass fraction). The responses to light, nutrients and water agreed with the (qualitative) prediction of the ‘functional equilibrium’ theory. The notable exception was atmospheric CO2, which did not affect allocation when the concentration was doubled. Third, we analysed the quantitative importance of the changes in allocation compared to changes in other growth parameters, such as unit leaf rate (the net difference between carbon gain and carbon losses per unit time and leaf area), and specific leaf area (leaf area: leaf biomass). The effects of light, CO2 and water on leaf mass fractions were small compared to their effects on relative growth rate. The effects of nutrients, however, were large, suggesting that only in the case of nutrients, biomass allocation is a major factor in the response of plants to limiting resource supply.