A Patchy Growth via Successive and Simultaneous Cambia: Key to Success of the Most Widespread Mangrove Species Avicennia marina?

BACKGROUND AND AIMS: Secondary growth via successive cambia has been intriguing researchers for decades. Insight into the mechanism of growth layer formation is, however, limited to the cellular level. The present study aims to clarify secondary growth via successive cambia in the mangrove species Avicennia marina on a macroscopic level, addressing the formation of the growth layer network as a whole. In addition, previously suggested effects of salinity on growth layer formation were reconsidered. METHODS: A 1-year cambial marking experiment was performed on 80 trees from eight sites in two mangrove forests in Kenya. Environmental (soil water salinity and nutrients, soil texture, inundation frequency) and tree characteristics (diameter, height, leaf area index) were recorded for each site. Both groups of variables were analysed in relation to annual number of growth layers, annual radial increment and average growth layer width of stem discs. KEY RESULTS: Between trees of the same site, the number of growth layers formed during the 1-year study period varied from only part of a growth layer up to four growth layers, and was highly correlated to the corresponding radial increment (0-5 mm year(-1)), even along the different sides of asymmetric stem discs. The radial increment was unrelated to salinity, but the growth layer width decreased with increasing salinity and decreasing tree height. CONCLUSIONS: A patchy growth mechanism was proposed, with an optimal growth at distinct moments in time at different positions around the stem circumference. This strategy creates the opportunity to form several growth layers simultaneously, as observed in 14 % of the studied trees, which may optimize tree growth under favourable conditions. Strong evidence was provided for a mainly endogenous trigger controlling cambium differentiation, with an additional influence of current environmental conditions in a trade-off between hydraulic efficiency and mechanical stability.     

Wood and bark anatomy of the African species of Gnetum

Quantitative and qualitative data are given for the two African species of Gnetum ( Gnetum section Gnetum subsection Micrognemones ). These species are lianoid and lack the fibre-tracheids of G. gnemon but have about the same vessel element and tracheid length as in that species. Vessel diameter is related to stem age and organography. Tori are clearly present in tracheary elements of the African Gnetum species, a first report for the genus. In these two species, origin of the lateral meristem, which produces vascular tissue and cambia, can be traced directly and indirecdy to cortical parenchyma. A second kind of meristematic action, newly reported for Gnetum, is produced by proliferation of axial parenchyma, fragmenting secondary xylem. Both presence of tori and site of origin of lateral meristematic activity in Gnetum contrast with corresponding conditions in Welwitschia.     

Wood and bark anatomy of Gnetutn gnemon L.

Quantitative and qualitative data are presented for seven collections representing two varieties (unlike in habit) of Gnetum gnemon. Tracheids are present, but abundant and intermixed with them are septate fibre-tracheids rich in starch. Axial parenchyma has been reported only once previously for the species. Axial parenchyma is in strands of 4–10 cells, is rich in starch, is primarily vasicentric (paratracheal) in distribution, less commonly diffuse. About equally common are simple and compound perforation plates; the latter are composed of from two to about ten bordered foraminate perforations, the shape of which may be altered by crowding or coalescence, but is clearly still foraminate. Lateral walls of vessels bear pits that are vestured around pit cavities, not facing the pit membrane. Rays are composed mostly of procumbent cells; the tangential walls bear bordered pits. Crystals, present in ray cells and (rarely) axial parenchyma vary widely in size. Crystalliferous sclereids with layered walls, starch-rich parenchyma, and gelatinous secondary phloem fibres are the main components of bark. Early stages in origin of successive vascular cambia in bark are newly described. When representative conditions are derived from study of large numbers of slides, the classical view that Gnetum vessels are unlike those of angiosperms is supported. Features of Gnetum gnemon wood are discussed in the light of ecology and conductive physiology.     

A mathematical framework for modelling cambial surface evolution using a level set method

Background and Aims

During their lifetime, tree stems take a series of successive nested shapes. Individual tree growth models traditionally focus on apical growth and architecture. However, cambial growth, which is distributed over a surface layer wrapping the whole organism, equally contributes to plant form and function. This study aims at providing a framework to simulate how organism shape evolves as a result of a secondary growth process that occurs at the cellular scale.

Methods

The development of the vascular cambium is modelled as an expanding surface using the level set method. The surface consists of multiple compartments following distinct expansion rules. Growth behaviour can be formulated as a mathematical function of surface state variables and independent variables to describe biological processes.

Key Results

The model was coupled to an architectural model and to a forest stand model to simulate cambium dynamics and wood formation at the scale of the organism. The model is able to simulate competition between cambia, surface irregularities and local features. Predicting the shapes associated with arbitrarily complex growth functions does not add complexity to the numerical method itself.

Conclusions

Despite their slenderness, it is sometimes useful to conceive of trees as expanding surfaces. The proposed mathematical framework provides a way to integrate through time and space the biological and physical mechanisms underlying cambium activity. It can be used either to test growth hypotheses or to generate detailed maps of wood internal structure.     

Stem and leaf anatomy of the arborescent Cucurbitaceae Dendrosicyos socotrana with comments on the evolution of pachycauls from lianas

Stem and leaf anatomy of Dendrosicyos socotrana, the only arborescent Cucurbitaceae, are examined for correlations with life form and ecology and are used to test hypotheses regarding features adaptive in scandent plants. The stem consists mainly of ray and conjunctive parenchyma with small strands of xylem forming an anastomosing net throughout the trunk. Xylem strands bear vascular cambia that produce secondary phloem, representing the first report of successive cambia in Cucurbitales. Some features characteristic of lianas, such as very wide vessel elements with thick walls, are absent from Dendrosicyos. Other features, such as very wide rays and abundant axial parenchyma, are present in both Dendrosicyos and lianas but appear to serve differing roles in these different life forms. It is suggested that lianas have numerous features that are readily co-opted in the evolution of pachycaul trees and that the evolution of pachycauls from lianas has happened repeatedly in the core eudicots.