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1.  Cambial Growth Season of Brevi-Deciduous Brachystegia spiciformis Trees from South Central Africa Restricted to Less than Four Months 
PLoS ONE  2012;7(10):e47364.
We investigate cambial growth periodicity in Brachystegia spiciformis, a dominant tree species in the seasonally dry miombo woodland of southern Africa. To better understand how the brevi-deciduous (experiencing a short, drought-induced leaf fall period) leaf phenology of this species can be linked to a distinct period of cambial activity, we applied a bi-weekly pinning to six trees in western Zambia over the course of one year. Our results show that the onset and end of cambial growth was synchronous between trees, but was not concurrent with the onset and end of the rainy season. The relatively short (three to four months maximum) cambial growth season corresponded to the core of the rainy season, when 75% of the annual precipitation fell, and to the period when the trees were at full photosynthetic capacity. Tree-ring studies of this species have found a significant relationship between annual tree growth and precipitation, but we did not observe such a correlation at intra-annual resolution in this study. Furthermore, a substantial rainfall event occurring after the end of the cambial growth season did not induce xylem initiation or false ring formation. Low sample replication should be taken into account when interpreting the results of this study, but our findings can be used to refine the carbon allocation component of process-based terrestrial ecosystem models and can thus contribute to a more detailed estimation of the role of the miombo woodland in the terrestrial carbon cycle. Furthermore, we provide a physiological foundation for the use of Brachystegia spiciformis tree-ring records in paleoclimate research.
PMCID: PMC3468463  PMID: 23071794
2.  A Patchy Growth via Successive and Simultaneous Cambia: Key to Success of the Most Widespread Mangrove Species Avicennia marina? 
Annals of Botany  2007;101(1):49-58.
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.
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.
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.
PMCID: PMC2701843  PMID: 18006508
Avicenia marina; cambial marking; mangrove; phloem; salinity; secondary growth; successive cambia; xylem
3.  Comparative Anatomy of Intervessel Pits in Two Mangrove Species Growing Along a Natural Salinity Gradient in Gazi Bay, Kenya 
Annals of Botany  2007;100(2):271-281.
Background and Aims
According to the air-seeding hypothesis, embolism vulnerability in xylem elements is linked directly to bordered pit structure and functioning. To elucidate the adaptive potential of intervessel pits towards fluctuating environmental conditions, two mangrove species with a distinct ecological distribution growing along a natural salinity gradient were investigated.
Scanning and transmission electron microscopic observations were conducted to obtain qualitative and quantitative characteristics of alternate intervessel pits in A. marina and scalariform intervessel pits in Rhizophora mucronata. Wood samples from three to six trees were collected at seven and five sites for A. marina and R. mucronata, respectively, with considerable differences between sites in soil water salinity.
Key Results
Vestured pits without visible pores in the pit membrane were observed in A. marina, the mangrove species with the widest geographical distribution on global as well as local scale. Their thick pit membranes (on average 370 nm) and minute pit apertures may contribute to reduced vulnerability to cavitation of this highly salt-tolerant species. The smaller ecological distribution of R. mucronata was in accordance with wide pit apertures and a slightly higher pitfield fraction (67 % vs. 60 % in A. marina). Nonetheless, its outer pit apertures were observed to be funnel-shaped shielding non-porous pit membranes. No trends in intervessel pit size were observed with increasing soil water salinity of the site.
The contrasting ecological distribution of two mangrove species was reflected in the geometry and pit membrane characteristics of their intervessel pits. Within species, intervessel pit size seemed to be independent of spatial variations in environmental conditions and was only weakly correlated with vessel diameter. Further research on pit formation and function has to clarify the large variations in intervessel pit size within trees and even within single vessels.
PMCID: PMC2735318  PMID: 17565970
Rhizophora mucronata; Avicennia marina; intervessel pits; salinity; Kenya; pit membrane; vestures; ecological wood anatomy; cavitation vulnerability; xylem; field-emission SEM; TEM
4.  Influence of a Salinity Gradient on the Vessel Characters of the Mangrove Species Rhizophora mucronata 
Annals of Botany  2006;98(6):1321-1330.
• Background and Aims Although mangroves have been extensively studied, little is known about their ecological wood anatomy. This investigation examined the potential use of vessel density as a proxy for soil water salinity in the mangrove species Rhizophora mucronata (Rhizophoraceae) from Kenya.
• Methods In a time-standardized approach, 50 wood discs from trees growing in six salinity categories were investigated. Vessel densities, and tangential and radial diameters of rainy and dry season wood of one distinct year, at three positions on the stem discs, were measured. A repeated-measures ANOVA with the prevailing salinity was performed.
• Key Results Vessel density showed a significant increase with salinity, supporting its use as a prospective measure of salinity. Interestingly, the negative salinity response of the radial diameter of vessels was less striking, and tangential diameter was constant under the varying environmental conditions. An effect of age or growth rate or the presence of vessel dimorphism could be excluded as the cause of the absence of any ecological trend.
• Conclusions The clear trend in vessel density with salinity, together with the absence of a growth rate and age effect, validates the potential of vessel density as an environmental proxy. However, it can only be used as a relative measure of salinity given that other environmental variables such as inundation frequency have an additional influence on vessel density. With view to a reliable, absolute proxy, future research should focus on finding wood anatomical features correlated exclusively with soil water salinity or inundation frequency. The plasticity in vessel density with differing salinity suggests a role in the establishment of a safe water transport system. To confirm this hypothesis, the role of inter-vessel pits, their relationship to the rather constant vessel diameter and the underlying physiology and cell biology needs to be examined.
PMCID: PMC2803579  PMID: 17062590
Rhizophora mucronata; mangrove; ecological wood anatomy; vessel density; vessel diameter; proxy; salinity; inundation frequency; Kenya; hydraulic architecture
5.  Growth Rings, Growth Ring Formation and Age Determination in the Mangrove Rhizophora mucronata 
Annals of Botany  2004;94(1):59-66.
• Background and Aims The mangrove Rhizophora mucronata has previously been reported to lack annual growth rings, thus barring it from dendrochronological studies. In this study the reported absence of the growth rings was reconsidered and the periodic nature of light and dark brown layers visible on polished stem discs investigated. In addition, the formation of these layers in relation to prevailing environmental conditions, as well as their potential for age determination of the trees, was studied.
• Methods Trees of known age were collected and a 2·5‐year cambial marking experiment was conducted to determine the periodic nature of the visible growth layers.
• Key Results Annual indistinct growth rings were detected in R. mucronata and are defined by a low vessel density earlywood and a high vessel density latewood. The formation of these growth rings and their periodic nature was independent from site‐specific environmental conditions in two forests along the Kenyan coast. However, the periodic nature of the rings was seriously affected by slow growth rates, allowing accurate age determination only in trees with radial growth rates above 0·5 mm year–1. The onset of the formation of the low vessel density wood coincided with the onset of the long rainy season (April–May) and continues until the end of the short rainy season (November). The high vessel density wood is formed during the dry season (December–March). Age determination of the largest trees collected in the two studied forests revealed the relatively young age of these trees (±100 years).
• Conclusions This study reports, for the first time, the presence of annual growth rings in the mangrove R. mucronata, which offers further potential for dendrochronological and silvicultural applications.
PMCID: PMC4242371  PMID: 15145790
Age determination; annual growth rings; cambial marking; dendrochronology; East Africa; mangroves; Rhizophora mucronata; wood anatomy

Results 1-5 (5)