We investigated scaling of physiological parameters between age classes of Quercus rubra by combining in situ field measurements with an experimental approach. In the in situ field study, we investigated changes in drought response with age in seedlings, juveniles, and mature trees of Q. rubra. Throughout the particularly dry summer of 1995 and the unusually wet summer of 1996 in New England, we measured water potential of leaves (Ψ(Leaf)) and gas exchange of plants at three sites at the Harvard Forest in Petersham, Massachusetts. In order to determine what fraction of the measured differences in gas exchange between seedlings and mature trees was due to environment versus ontogeny, an experiment was conducted in which seedlings were grown under light and soil moisture regimes simulating the environment of mature trees. The photosynthetic capacity of mature trees was three-fold greater than that of seedlings during the wet year, and six-fold greater during the drought year. The seedling experiment demonstrated that the difference in photosynthetic capacity between seedlings and mature trees is comprised equally of an environmental component (50%) and an ontogenetic component (50%) in the absence of water limitation. Photosynthesis was depressed more severely in seedlings than in mature trees in the drought year relative to the wet year, while juveniles showed an intermediate response. Throughout the drought, the predawn leaf water potential (Ψ(PD)) of seedlings became increasingly negative (-0.4 to -1.6 MPa), while that of mature trees became only slightly more negative (-0.2 to -0.5 MPa). Again, juveniles showed an intermediate response (-0.25 to -0.8 MPa). During the wet summer of 1996, however, there was no difference in Ψ(PD) between seedlings, juveniles and mature trees. During the dry summer of 1995, seedlings were more responsive to a major rain event than mature trees in terms of Ψ(Leaf), suggesting that the two age classes depend on different water sources. In all age classes, instantaneous measurements of intrinsic water use efficiency (WUE(i)), defined as C assimilation rate divided by stomatal conductance, increased as the drought progressed, and all age classes had higher WUE(i) during the drought year than in the wet year. Mature trees, however, showed a greater ability to increase their WUE(i) in response to drought. Integrated measurements of WUE from C isotope discrimination (δ) of leaves indicated higher WUE in mature trees than juveniles and seedlings. Differences between years, however, could not be distinguished, probably due to the strong bias in C isotope fractionation at the time of leaf production, which occurred prior to the onset of drought conditions in 1995. From this study, we arrive at two main conclusions: 1. Different age classes of Q. rubra use different strategies for responding to drought. Seedlings resist drought by closing stomata early in the day at the expense of C uptake; mature trees avoid drought conditions by accessing deeper water reserves and adjusting WUE. 2. Only through studies which separate environmental differences from ontogenetic differences can parameters measured on seedlings be scaled to mature trees.
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Acknowledgements The authors thank S. Chun for assistance in the field, S. Bassow for technical assistance, J. Montoya and N. van der Merwe for assistance with the stable isotope analyses, and E. Macklin for assistance with the statistical analyses. K. Newkirk, J. Humphries, and J. Ehleringer provided unpublished data sets from the Harvard Forest. D. Ackerly, G. Berntson, P. Wayne, K. Stinson, E. Macklin, D. Schrag and an anonymous reviewer are gratefully acknowledged for helpful discussions and comments on the manuscript. This work was supported by the Mellon Foundation and an NSF grant to the Harvard Forest LTER (DEB 9411795).
- Drought response
- Quercus rubra
- Water use efficiency