Using growth analysis to interpret competition between a C3 and a C4 annual under ambient and elevated CO2

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Abstract
Detailed growth analysis in conjunction with information on leaf display and nitrogen uptake was used to interpret competition between Abutilon theophrasti, a C3 annual, and Amaranthus retroflexus, a C4 annual, under ambient (350 μl l-1) and two levels of elevated (500 and 700 μl l-1) CO2. Plants were grown both individually and in competition with each other. Competition caused a reduction in growth in both species, but for different reasons. In Abutilon, decreases in leaf area ratio (LAR) were responsible, whereas decreased unit leaf rate (ULR) was involved in the case of Amaranthus. Mean canopy height was lower in Amaranthus than Abutilon which may explain the low ULR of Amaranthus in competition. The decrease in LAR of Abutilon was associated with an increase in root/shoot ratio implying that Abutilon was limited by competition for below ground resources. The root/shoot ratio of Amaranthus actually decreased with competition, and Amaranthus had a much higher rate of nitrogen uptake per unit of root than did Abutilon. These latter results suggest that Amaranthus was better able to compete for below ground resources than Abutilon. Although the growth of both species was reduced by competition, generally speaking, the growth of Amaranthus was reduced to a greater extent than that of Abutilon. Regression analysis suggests that the success of Abutilon in competition was due to its larger starting capital (seed size) which gave it an early advantage over Amaranthus. Elevated CO2 had a positive effect upon biomass in Amaranthus, and to a lesser extent, Abutilon. These effects were limited to the early part of the experiment in the case of the individually grown plants, however. Only Amaranthus exhibited a significant increase in relative growth rate (RGR). In spite of the transitory effect of CO2 upon size in individually grown plants, level of CO2 did effect final biomass of competitively grown plants. Abutilon grown in competition with Amaranthus had a greater final biomass than Amaranthus at ambient CO2 levels, but this difference disappeared to a large extent at elevated CO2. The high RGR of Amaranthus at elevated CO2 levels allowed it to overcome the difference in initial size between the two species.