Abstract
Fruiting is typically considered to massively burden the seasonal carbon budget of trees. The cost of reproduction has therefore been suggested as a proximate factor explaining observed mast-fruiting patterns. Here, we used a large-scale, continuous 13C labeling of mature, deciduous trees in a temperate Swiss forest to investigate to what extent fruit formation in three species with masting reproduction behavior (Carpinus betulus, Fagus sylvatica, Quercus petraea) relies on the import of stored carbon reserves. Using a free-air CO2 enrichment system, we exposed trees to 13C-depleted CO2 during 8 consecutive years. By the end of this experiment, carbon reserve pools had significantly lower δ13C values compared to control trees. δ13C analysis of new biomass during the first season after termination of the CO2 enrichment allowed us to distinguish the sources of built-in carbon (old carbon reserves vs. current assimilates). Flowers and expanding leaves carried a significant 13C label from old carbon stores. In contrast, fruits and vegetative infructescence tissues were exclusively produced from current, unlabeled photoassimilates in all three species, including F. sylvatica, which had a strong masting season. Analyses of δ13C in purified starch from xylem of fruit-bearing shoots revealed a complete turn-over of starch during the season, likely due to its usage for bud break. This study is the first to directly demonstrate that fruiting is independent from old carbon reserves in masting trees, with significant implications for mechanistic models that explain mast seeding.





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Acknowledgments
The authors would like to thank Catharina Lötscher for starch extraction and 13C analyses, Erwin Amstutz for operating the crane, Daisuke Kabeya for helping in the field and his valuable discussions on tree reproduction, and Olivier Bignucolo, Susanna Riedl and Gabrielle Schaer for litter trap collection and fruit biomass analyses. We further thank the two anonymous referees for their valuable comments and their helpful suggestions to improve the text. This study was partially supported by grants from the Japanese Society for the Promotion of Science (No. 18580155, No. 21380103 to Q.H.) and a research fellowship to Q.H. from the Co-operative OECD Research Program. The CO2 enrichment and crane infrastructure at the SCC site were funded by grants to C.K. from the Swiss National Science Foundation (SNF, No. 3100-059769.99, No. 3100-067775.02) and NCCR-Climate (No. 5005-65755). During the time of writing this manuscript, G.H. received funding from European Research Council (ERC) grant No. 233399 (project ‘TREELIM’ to C.K.).
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Communicated by Russell Monson.
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Hoch, G., Siegwolf, R.T.W., Keel, S.G. et al. Fruit production in three masting tree species does not rely on stored carbon reserves. Oecologia 171, 653–662 (2013). https://doi.org/10.1007/s00442-012-2579-2
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DOI: https://doi.org/10.1007/s00442-012-2579-2
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