title: Flood-Ring Formation and Root Development in Response to Experimental Flooding of Young Quercus robur Trees.
abstract: Spring flooding in riparian forests can cause significant reductions in earlywood-vessel size in submerged stem parts of ring-porous tree species, leading to the presence of 'flood rings' that can be used as a proxy to reconstruct past flooding events, potentially over millennia. The mechanism of flood-ring formation and the relation with timing and duration of flooding are still to be elucidated. In this study, we experimentally flooded 4-year-old Quercus robur trees at three spring phenophases (late bud dormancy, budswell, and internode expansion) and over different flooding durations (2, 4, and 6 weeks) to a stem height of 50 cm. The effect of flooding on root and vessel development was assessed immediately after the flooding treatment and at the end of the growing season. Ring width and earlywood-vessel size and density were measured at 25- and 75-cm stem height and collapsed vessels were recorded. Stem flooding inhibited earlywood-vessel development in flooded stem parts. In addition, flooding upon budswell and internode expansion led to collapsed earlywood vessels below the water level. At the end of the growing season, mean earlywood-vessel size in the flooded stem parts (upon budswell and internode expansion) was always reduced by approximately 50% compared to non-flooded stem parts and 55% compared to control trees. This reduction was already present 2 weeks after flooding and occurred independent of flooding duration. Stem and root flooding were associated with significant root dieback after 4 and 6 weeks and mean radial growth was always reduced with increasing flooding duration. By comparing stem and root flooding, we conclude that flood rings only occur after stem flooding. As earlywood-vessel development was hampered during flooding, a considerable number of narrow earlywood vessels present later in the season, must have been formed after the actual flooding events. Our study indicates that root dieback, together with strongly reduced hydraulic conductivity due to anomalously narrow earlywood vessels in flooded stem parts, contribute to reduced radial growth after flooding events. Our findings support the value of flood rings to reconstruct spring flooding events that occurred prior to instrumental flood records.
authors.affiliation: Forest Ecology and Forest Management Group, Wageningen University and Research CentreWageningen, Netherlands; Alterra, Wageningen University and Research CentreWageningen, Netherlands.
authors.forename: Paul
authors.initials: P
authors.lastname: Copini
authors.affiliation: Forest Ecology and Forest Management Group, Wageningen University and Research Centre Wageningen, Netherlands.
authors.forename: Jan
authors.initials: J
authors.lastname: den Ouden
authors.affiliation: Laboratory of Wood Biology and Xylarium, Royal Museum for Central AfricaTervuren, Belgium; Laboratory of Plant Biology and Nature Management, Vrije Universiteit BrusselBrussels, Belgium.
authors.forename: Elisabeth M R
authors.initials: EM
authors.lastname: Robert
authors.affiliation: Centre for Forest Interdisciplinary Research, Department of Biology, The University of Winnipeg Winnipeg, Canada.
authors.forename: Jacques C
authors.initials: JC
authors.lastname: Tardif
authors.affiliation: Forest Ecology and Forest Management Group, Wageningen University and Research Centre Wageningen, Netherlands.
authors.forename: Walter A
authors.initials: WA
authors.lastname: Loesberg
authors.affiliation: Forest Ecology and Forest Management Group, Wageningen University and Research Centre Wageningen, Netherlands.
authors.forename: Leo
authors.initials: L
authors.lastname: Goudzwaard
authors.affiliation: Forest Ecology and Forest Management Group, Wageningen University and Research Centre Wageningen, Netherlands.
authors.forename: Ute
authors.initials: U
authors.lastname: Sass-Klaassen
keywords: Quercus robur; flooding; hypoxia; leaf phenology; pedunculate oak; root development; vessel development