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Detailed and four dimensional - the three spatial dimensions and time - observations of these xylem and phloem networks are lacking, though highly necessary to understand Avicennia's growth pattern and the secondary growth of trees with successive cambia in general.ДЇrom a study in two dimensions, the nature of secondary growth in Avicennia marina has been suggested to be patch-like, with active growth displacing around the stem circumference with time. 1c), ,, ,, , and form an intricate three-dimensional network instead of separate concentric cylinders. Xylem and phloem tissue in Avicennia are found to be non-concentric ( Fig. The genus Avicennia is the only mangrove tree genus showing the developmental oddity of secondary growth by successive cambia. Special forms of secondary wood formation are typical for lianas and vines, ,, ,, though the phenomenon of successive cambia is also found in a select group of herb, shrub and tree species, ,, ,, coming from different phylogenetic clades.

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The complexity of the internal structure can be expressed by the sum of the locations of free ending and branched phloem tissue (f) per stem disc surface area. On the micro-CT-images (g-k) the wide light grey bands are xylem tissue while the dark small strings are phloem tissue. Each section is 150 Вµm apart from the previous.

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This is shown by the white arrowheads in the four serial micro-CT-scan images of an outside zone of an Avicennia marina stem disc, produced by a SkyScan 1172 scanner (g-k). The connections between xylem patches changed rapidly with vertical distance. It can be seen that Avicennia had a complex network of xylem patches that fused at certain heights of the tree and joined different patches at other parts of the stem. Only a small part of the xylem tissue has been visualised. The spatial structure of Avicennia is depicted by smoothed surface images after three-dimensional reconstruction in Reconstruct (d,e).

Schematic view of a stem disc from (a) a tree with only one cambium, (b) a tree with successive cambia organised in concentric cylinders, giving rise to a stem disc with concentric circles of xylem tissue, phloem tissue and cambium circles and (c) Avicennia, having a reticulate organisation of its cambia and transport structure. Spatial structure of the xylem and phloem tissue in Avicennia in comparison to other trees. Herewith the authors confirm that this does not alter their adherence to all the PLoS ONE policies on sharing data and material, as detailed in the PLoS Policy on Declaration and Evaluation of Competing Interests.ДЇigure 1. The company in no way obstructs publication and sharing of the data used for this publication. The input of the company was of a purely academic and technical nature. There is no other affiliation with Skyscan NV than scientific contacts. Published with the financial support of the Universitaire Stichting вЂђ BelgiГ«, Egmontstraat 11, BвЂђ1000 Brussels, Belgium.Д¬ompeting interests: Elke Van de Casteele was, at the time the study was conducted, employed by Skyscan NV, Kontich, Belgium, which offered free micro-CT scanning. The funders had no role in study design, content of the study, decision to publish or determination of the conclusions. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.ДЇunding: The study was financially supported by the Institute for the Promotion of Innovation through Science and Technology in Flanders (IWT-Vlaanderen), the National Fund for Scientific Research (FWO, Belgium), the King Leopold III Fund for Nature Exploration and Conservation, Belgium, and the AFRODENDRO project of the Royal Museum for Central Africa (RMCA), Tervuren. Received: SeptemAccepted: DecemPublished: January 31, 2011Д¬opyright: В© 2011 Robert et al. (2011) Successive Cambia: A Developmental Oddity or an Adaptive Structure? PLoS ONE 6(1):Д®ditor: Wendy Peer, Purdue University, United States of America Citation: Robert EMR, Schmitz N, Boeren I, Driessens T, Herremans K, De Mey J, et al.