Is Cork Growth a Reliable Proxy for Stem Diameter Growth in Cork Oak (Quercus suber L.)? Implications for Forest Management under Climate Change in Mediterranean Regions

Cork-ring widths have been extensively used in dendroecological studies assessing the relationship between cork growth patterns and climate (precipitation and temperature). Generally, cork growth is assumed as a proxy for stem diameter growth to address cork oak (Quercus suber L.) growth sensitivity to climate and cork yield modeling. Cork growth represents a large part of stem radial increment in this species due to the enhanced activity of phellogen when compared to the cambium activity; thus, similar inter-annual variations of cork-ring widths and tree diameter growth might be expected. However, so far, the influence of rainfall and temperature on stem diameter growth has scarcely been addressed; moreover, it is still not clear whether tree size relates, and in what way, to the variations in radial growth of cork and stem diameter and whether these reflect (proportional) quantitative variations in stem basal area growth. In this study, we computed the annual growth of cork and of stem diameter at breast-height in data series of 47 trees, from 2000 to 2012, corresponding to a full cork production cycle. Results showed a tight link between cork-ring width and stem diameter growth indices. However, while cork growth strongly correlated with climate conditions in autumn–winter prior to the growing season, stem diameter growth correlated with climate conditions of the current growing season, and, more importantly, it was tree size-related. The extrapolation from cork-ring increments to stem basal area growth is likely to progressively underestimate tree growth and biomass increment in larger cork oaks and to further bias it due to climate change effects in the Mediterranean region.

Transcriptomic analysis of cork during seasonal growth highlights regulatory and developmental processes from phellogen to phellem formation

The phellogen or cork cambium stem cells that divide periclinally and outwardly specify phellem or cork. Despite the vital importance of phellem in protecting the radially-growing plant organs and wounded tissues, practically only the suberin biosynthetic process has been studied molecularly so far. Since cork oak (Quercus suber) phellogen is seasonally activated and its proliferation and specification to phellem cells is a continuous developmental process, the differentially expressed genes during the cork seasonal growth served us to identify molecular processes embracing from phellogen to mature differentiated phellem cell. At the beginning of cork growth (April), cell cycle regulation, meristem proliferation and maintenance and processes triggering cell differentiation were upregulated, showing an enrichment of phellogenic cells from which phellem cells are specified. Instead, at maximum (June) and advanced (July) cork growth, metabolic processes paralleling the phellem cell chemical composition, such as the biosynthesis of suberin, lignin, triterpenes and soluble aromatic compounds, were upregulated. Particularly in July, polysaccharides- and lignin-related secondary cell wall processes presented a maximal expression, indicating a cell wall reinforcement in the later stages of cork formation, presumably related with the initiation of latecork development. The putative function of relevant genes identified are discussed in the context of phellem ontogeny.

Drivers for Annual Cork Growth under Two Understory Management Alternatives on a Podzolic Cork Oak Stand

Understory management practices and stand density characteristics allow one to distinguish a cork oak traditional silvopastoral system (known as a montado) from a cork oak forest system. Although understanding the manner in which different management practices affect cork growth is imperative, there are still only a few outputs from experimental research that contribute to this knowledge. The effect of potential drivers on annual cork growth was analyzed using a linear mixed model approach. Two dimensions of drivers were considered: intraspecific competition, assessed by tree level distance-dependent indices; and interspecific competition, assessed by variables characterizing understory management. The present dataset was collected from an experimental trial established on a cork oak stand in Podzolic soil on the Tagus river basin, covering two different cork growth cycles over the period from 2003 to 2015. The adjusted models considered two understory management alternatives: spontaneous shrubs maintenance and forage application. In both models, annual precipitation displayed a positive effect on annual cork growth, as expected. However, no significant effect of intraspecific competition was found. Additionally, there was a positive effect on annual cork growth associated with the spontaneous shrubs growth and a negative effect associated with lupine presence; both effects linked to different cork ring ages’ thresholds. The study main contributions are the following: (i) the introduction of the interaction between cork growth cycle stage and understory management practices, only possible with cork sample collections from different cork rotation cycles; (ii) the finding that there was no significant effect of intraspecific competition on cork growth.