Pith: a marker of primary growth in Picea abies (L.) Karst.
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The primary growth of trees may be studied either by following their development over time, which is costly and requires long-term monitoring, or by a posteriori growth analyses. Trees in temperate forests show rhythmic growth, which is characterised by morphological or anatomical markers. The study described here focuses on the pith of trees as an internal marker for the retrospective analysis of primary growth. Changes in pith size and density were quantified along a stem of Norway spruce [Picea abies (L.) Karst.] with high spatial resolution. The results showed that pith would appear to be a reliable marker of the annual growth rhythm even in the presence of polycyclism. Annual shoot limits were characterised by reductions in pith size associated, at the same time, with increases in pith density. In addition, pith size may provide information about tree ontogeny. The start of the competition with neighbouring trees was very likely responsible for an overall decrease in pith size. Regarding high frequency variations, pith size appeared to be less sensitive to local environmental fluctuations like climate than other studied variables such as annual shoot length and annual ring width. Finally, X-ray computed tomography proved to be a very promising method for the non-destructive detection of annual shoot limits in stems based on longitudinal changes in pith density, as demonstrated in a log of Norway spruce.
KeywordsAnnual shoot Rhythmic growth Pith density Pith size Norway spruce
The authors wish to thank Pierre Gelhaye, LERFoB Nancy, for his advice and assistance with sampling in the field and at the laboratory, and for the 2D X-ray measurements. Thanks to Charline Freyburger, LERFoB Nancy, for the measurements made by means of X-ray computed tomography. Thanks also to Patrick Heuret for helpful discussions about tree architecture. AMAP (Botany and Computational Plant Architecture) is a joint research unit which associates CIRAD (UMR51), CNRS (UMR5120), INRA (UMR931), IRD (R123), and Montpellier 2 University (UM27); http://amap.cirad.fr/.
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