The balanced-growth hypothesis and the allometry of leaf and root biomass allocation

1. Many ecological models of plant growth assume balanced growth: that biomass is allocated preferentially to leaves or roots to increase capture of the limiting external resource. An alternative explanation is based on nonlinear (allometric) allocation as a function of plant size. The objective of this study was to test between these two alternative explanations. 2. A total of 1150 plants from 22 different herbaceous species were grown in hydroponic sand culture in factorial combinations of high (1100 mumol m(-2) s(-1)) and low (200 mumol m(-2) s(-1) PAR) irradiance crossed with a full-strength and a 1/6 dilution of Hoagland's hydroponic solution. Plants were harvested at 15, 20, 25, 30 and 35 days postgermination, and dry mass was determined for leaf and root components. These data were used to test the hypotheses of balanced growth and of allometric allocation. 3. Both irradiance and nutrient supply affected the slope and intercept of the root : shoot allometry, contrary to the allometric hypothesis but in agreement with the hypothesis of balanced growth; decreased nutrient supply increased allocation to roots; and decreased irradiance increased allocation to leaves. 4. Plants allocated relatively more biomass to roots than to leaves as plants grew larger. In order for the balanced-growth hypothesis to be correct, the net rate of nutrient uptake per unit root mass must have been decreasing relative to the net rate of carbon gain per unit leaf mass. 5. We suggest two reasons why this might be the case: (i) older roots decreased their efficiency of nutrient uptake; and (ii) larger root systems more rapidly decreased the available nutrients between flushes of hydroponic solution. 6. These results support the notion of balanced growth that is found in many ecological models of plant growth.