Xylem and Phloem Formation Dynamics in Quercus ilex L. at a Dry Site in Southern Italy

Abstract

Quercus ilex L. dieback has been recently reported at numerous Mediterranean sites. Wood and phloem formation dynamics and tree-ring series of anatomical traits can be used to evaluate growth conditions of trees. We monitored cambial activity in Q. ilex trees growing at a site in southern Italy in order to assess how xylem and phloem production are affected by harsh seasonal climatic variation during a dry year. We followed xylogenesis by counting the number of cambial cells and detecting the occurrence of post-cambial cells throughout the year. As phloem did not show clear growth rings and boundaries between them, we followed the development of phloem fibres—their morphological traits during development and the distance from the cambium served as a reference point to evaluate the phloem production during the year. We detected a multimodal pattern in cambial activity, with wood production in three periods of the year and consequent formation of intra-annual density fluctuations (IADFs). The lowest production of xylem cells was observed in the dry late spring and summer period (likely due to the low water availability), while the highest occurred in autumn (the wettest period). Although we could not differentiate between early and late phloem, the analysis of phloem traits was useful to follow the dynamics of phloem production, which is generally difficult in Mediterranean tree species. We found cambial production of phloem throughout the year, even in the periods without xylem production. The results showed that if tree growth was constrained by environmental limitations, the ratio between xylem to phloem cells decreased and, in the most severely affected trees, more cells were formed preferentially in the phloem compared to xylem. We also briefly report the way in which to solve technical problems with tissue preparation due to extreme hardness and to the peculiar structure of Q. ilex wood and outer bark.