ALBERT

Notes: Four- to seven-year-old spruce trees (Picea abies) were exposed to three CO2 concentrations (280, 420 and 560 cm3 m−3) and three rates of wet N deposition (0, 30 and 90 kg ha−1 year−1) for 3 years in a simulated montane forest climate. Six trees from each of six clones were grown in competition in each of nine 100 × 70 × 36 cm model ecosystems with nutrient-poor natural forest soil. Stem dises were analysed using X-ray densitometry. The radial stem increment was not affected by [CO2] but increased with increasing rates of N deposition. Wood density was increased by [CO2], but decreased by N deposition. Wood-starch concentration increased, and wood nitrogen concentration decreased with increasing [CO2], but neither was affected by N deposition. The lignin concentration in wood was affected by neither [CO2] nor N deposition. Our results suggest that, under natural growth conditions, rising atmospheric [CO2] will not lead to enhanced radial stem growth of spruce, but atmospheric N deposition will, and in some regions is probably already doing so. Elevated [CO2], however, will lead to denser wood unless this effect is compensated by massive atmospheric N deposition. If can be speculated that greater wood density under elevated [CO2] may alter the mechanical properties of wood, and higher ratios of C/N and lignin/N in wood grown at elevated [CO2] may affect nutrient cycles of forest ecosystems.

Notes: Few of the most common assumptions used in models of responses of plants and ecosystems to elevated CO2 and climate warming have been tested under realistic life con-ditions. It is shown that some unexpected discrepancies between predictions and experimental findings exist, suggesting that a better empirical basis is required for predictions. The following ten suggestions may improve our potential to scale up from experimental scales to the real world.(1) Experiments should be timed to account for non-linearity in system responsiveness, asynchrony of responses and developmental differences. (2) By altering mineral nutrient supply, a wide range of CO2 responses can be ‘produced’, thus requiring realistic soil conditions. (3) Distinctions should be made between ‘doubling CO2 sup-ply’ and biologically effective degrees of CO2 enrichment. (4) Because of the non-linearity of plant responses to CO2, studies of at least three instead of two CO2 concentrations are necessary to describe future trends adequately. (5) Edge effects, in particular unscreened side light, may lead to allometric anomalies, strongly constraining up-scaling to stand-scale CO2 responses. (6) Variables such as growth, yield, net primary production and C turnover are often confused with carbon pools, carbon sequestration or net ecosystem production. (7) Mono- and interspecific interactions between individuals may lead to completely unpredictable CO2 responses. (8) Experiments with seedlings benefit from the absence of prehistory effects but are likely to be irrelevant for the responses of larger trees which, on the other hand, may be constrained by carry-over effects. Tree ring research indicates immediate sensitivity of large trees to environmental changes, supporting their usefulness in short-term CO2-enrichment experiments. (9) In predicting temperature responses, acclimation deserves more attention. (10) The significance of developmental responses is largely under-represented in experimental research, although these responses may overrule many of the other effects of atmospheric change. Results of more realistic experiments which account for these problems will provide a better basis for modelling the future of the biosphere.

Notes: Of the many responses of plants to elevated CO2, accumulation of total non-structural carbohydrates (TNC in % dry weight) in leaves is one of the most consistent. Insufficient sink activity or transport capacity may explain this obvious disparity between CO2 assimilation and carbohydrate dissipation and structural investment. If transport capacity contributes to the problem, phloem loading may be the crucial step. It has been hypothesized that symplastic phloem loading is less efficient than apoplastic phloem loading, and hence plant species using the symplastic pathway and growing under high light and good water supply should accumulate more TNC at any given CO2 level, but particularly under elevated CO2. We tested this hypothesis by carrying out CO2 enrichment experiments with 28 plant species known to belong to groups of contrasting phloem-loading type. Under current ambient CO2 symplastic loaders were found to accumulate 36% TNC compared with only 19% in apoplastic loaders (P=0.0016). CO2 enrichment to 600 μmol mol−1 increased TNC in both groups by the same absolute amount, bringing the mean TNC level to 41% in symplastic loaders (compared to 25% in apoplastic loaders), which may be close to TNC saturation (coupled with chlornplast malfunction). Eight tree species, ranked as symplastic loaders by their minor vein companion cell configuration, showed TNC responses more similar to those of apoplastic herbaceous loaders. Similar results are obtained when TNC is expressed on a unit leaf area basis, since mean specific leaf areas of groups were not significantly different. We conclude that phloem loading has a surprisingly strong effect on leaf tissue composition, and thus may translate into alterations of food webs and ecosystem functioning, particularly under high CO2.