Variation of wood density and hydraulic properties of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) clones related to a heat and drought wave in France

Drought response of three Douglas-fir clones (slow, intermediate and fast growing) inhabiting two different climatic regimes in France was examined. We used the hydraulic conductivity and the percent loss of conductivity due to embolism from stems and branches as well as wood microdensity measurements to determine the role of genetics in the control of embolism in this species. Conductivity and tree-ring’s microdensity variables (ring width: RW, mean ring density: MRD, minimum ring density: MID and maximum ring density: MAD) were compared in growth rings in all three clones during a typical year (2002) vs. an extremely dry year (2003). A new method was developed in order to assess axial hydraulic specific conductivity (K_s) within tree rings. The results show that branches are more resistant to embolism than stems, and that there are significant differences in embolism resistance among the clones between the two sites. K_s varied between years, sites and clones but the site exerted the most significant effect. Lartimache (more humid site during 2003) trees showed substantially higher K_s than those inhabiting Chassenoix (strongly affected by the 2003 heat and drought wave site). Wood analysis showed a significant year effect for all ring variables and a significant clone effect for all ring variables except for MAD, while the site effect was significant for MRD and MAD. The existence of a genetic control for the study traits indicates that Douglas-fir populations introduced in France may have a selection potential to face extreme climate events like the 2003 heat and drought wave.     

What is hot in tree rings? The wood density of surviving Douglas-firs to the 2003 drought and heat wave

In a global warming scenario, drought and heat waves like the one that occurred in 2003 in Europe are expected to become more intense and frequent. This extreme climate event strongly affected the hydraulic balance in many forest tree species including Douglas-fir, with symptoms ranging from partial crown necrosis to death. We studied a French Douglas-fir provenance trial, strongly affected by the 2003 drought and heat wave. Using wood X-ray microdensity profiles as a record of cambium response to environmental variations, we compared mean stem density and growth between dead and alive neighbouring trees and several microdensity characters measured over 17 growth rings previous to 2003. Special attention has been given to the sampling layout in order to minimize the micro-environmental effects. At tree level, surviving trees have a significantly higher mean stem density. At tree ring level, they have a significantly higher mean ring density, maximum ring density, latewood density and, to a lesser extent, latewood proportion. No significant difference was found for diameter growth. These results could have direct implications in the way to study and predict possible acclimation and adaptation of forest trees to climate change process. Wood could be used for the selection of genotypes with a desirable plasticity and resistance to drought induced-cavitation.     

Genetic control of the tree-ring response of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) to the 2003 drought and heat-wave in France

The genetic control of tree ring growth in Douglas-fir in response to the drought and heat-wave that occurred in 2003 in Europe was studied with microdensity profiles in three clonal experiments located in three different French regions. The drought and heat-wave significantly affected Douglas-fir wood formation. The Chassenoix site (Northeast of Massif-Central) was more severely affected that the other two: the 2003 year-ring was narrower and less dense than in the other sites and than the previous (2002) and following (2004) year-rings in the same site: ring growth stopped earlier and latewood did not develop completely. The year-rings 2004 and 2002 were very similar in this site. There was a significant genetic control for all ring parameters in the three sites and during the three years. The heritability was highly variable between years and among sites, without any clear pattern in this variability, except in Chassenoix where it was slightly lower in the 2003 year-ring. Variables measuring the response of trees to the 2003 event, i.e., the difference in ring width between 2002 and 2003, or between 2003 and 2004, showed a very variable degree of genetic control, from very low to relatively high. Douglas-fir seemed plastic enough to acclimate to the drought and heat-wave and then to recover during 2004. Furthermore the level of heritability estimated demonstrates that Douglas-fir has an adaptive potential that could be useful for multi-generation long-term response.     

Influence of cambial age and climate on ring width and wood density in Pinus radiata families

? Context

The correlation between tree ring width and density and short-term climate fluctuations may be a useful tool for predicting response of wood formation process to long-term climate change.

? Aims

This study examined these correlations for different radiata pine genotypes and aimed at detecting potential genotype by climate interactions.

? Methods

Four data sets comprising ring width and density of half- and full-sib radiata pine families were used. Correlations with climate variables were examined, after the extraction of the effect of cambial age.

? Results

Cambial age explained the highest proportion of the ring to ring variation in all variables. Calendar year and year by family interaction explained a smaller but significant proportion of the variation. Rainfall had a positive correlation with ring width and, depending on test site, either a negative or positive correlation with ring density. Correlations between temperature during growing season and ring density were generally negative.

? Conclusion

Climate variables that influence ring width and wood density can be identified from ring profiles, after removing the cambial age effect. Families can be selected that consistently show desirable response to climate features expected to become prevalent as a result of climate change.     

Drought resistance of two hybrid Populus clones grown in a large-scale plantation

Poplar hybrids were grown with irrigation in a large-scale plantation to investigate the mechanisms underlying clonal differences in drought resistance. Beginning in spring 1992, Populus trichocarpa x P. deltoides (TD) and P. deltoides x P. nigra (DN) cuttings received 46, 76, or 137 cm year(-1) of irrigation to supplement the 18-20 cm of annual precipitation, and all trees received the same fertilization regime. Stem volume, assessed as the square of stem diameter at breast height times tree height (D(2)H), and water relations of the trees were studied from the end of their second growing season until the end of their fifth growing season. By the end of the second growing season, stem volume of Clone TD was 40-146% larger than that of Clone DN, but stem volume growth was independent of irrigation in excess of 46 cm year(-1) in both clones. During the third growing season, stem volume growth of both clones was limited by both the 46- and 76-cm irrigation treatments, so that by the end of the third growing season trees in the 46-cm irrigation treatment were only half the size of trees in the 137-cm irrigation treatment. These treatment differences were maintained through the fifth growing season. Although stem volumes of Clone TD trees in the 76- and 137-cm irrigation treatments were larger than the corresponding values for Clone DN trees at the end of the third growing season (1994), these clonal differences gradually decreased in subsequent years and were not detectable after 5 years, because stem volume relative growth rate of Clone DN was greater than that of Clone TD in all treatments. Although both clones exhibited similar predawn leaf water potentials, Clone DN typically maintained higher midday leaf water potentials, suggesting better stomatal control of water loss. Clonal and treatment differences in osmotic potential at full turgor were minimal and could not explain the clonal differences in drought resistance. Root density and root density to stem volume ratio increased more in response to moderate drought in Clone DN than in Clone TD, resulting in enhanced drought resistance (high stem volume growth rate under moderate drought conditions) and an increased capacity to withdraw water from the soil. We conclude that the greater drought resistance of Clone DN compared with Clone TD was the result of the maintenance of a more favorable water balance by stomatal regulation and greater carbon allocation to roots during the early stages of drought. However, the low root density to stem volume ratio in Clone DN growing in the 46-cm irrigation treatment suggests that severe water limitation restricted the preferential allocation of carbon to belowground tissues, so that both root and shoot growth were constrained by severe drought.     

乔木柳无性系木材基本朱内变异的研究

利用乔木柳12年生无性系对比试验林的材料，研究了13个柳树无性系木材基本密度株内径向和纵向的变异模式，并对密度选择的时期和取样位置进行了探讨。结果表明：（1）在1.3m高度，乔木柳无性系各年轮组木材戏向变异多数为“M”型曲线，即从木芯向外第、至第3年轮组（1－6a）的木材密度逐渐增加，第4年轮组（7－8a）略有下降，第5年轮组（9－10a）又有明显增加，至最外侧年轮（11－12a）木材密度则显著下降，根据各无性系的变异特点，分成3种类型，其中无性系221的变异较为独特，为典型的“N”型；（2）乔木柳无性系生长期密度变化的规律为：1－2年生时木材基本密度相对较低，3－4a以后密度相对稳定并接近无性系加权平均密度，3－4年生时木材基本密度与5－12年生时的测定值相关系数为0.789－0.985，均达到极显著相关水平（α＝0.01），5－12年生乔木柳无性系木材基本密度的变异为62.2％－97.0％与3－4年生时木材基本密度的值有关，3－4a作为5百木柳木材基本密度的选择年龄最为合适；（3）供试群体木材密度株内纵向变异模式大多也可用“M”型曲线加以描述，具典型的苴孔材绷向密度变异特征，根据各无性系的变异特点，分成4种变异类型；树干高1.3，3.6和7.2m处的木材基本密度与无性系平均密度的相关系数达到显著和极显著水平，可以在树干1.3m处取样以推测无性系平均值；（4）在供试群体中，有的无性系木材基本密度纵向和径向变异的幅度均很小，木材较为均一，如苏柳791，表明在乔木柳杂种后代中选择木材均一的柳树新品种是可能的。     

False ring formation in teak (Tectona grandis L.f.) and the influence of environmental factors

Seedling anatomy and cross sectional discs of 8- and 12-year-old (juvenile) teak trees were examined for tree ring analysis and determination of environmental factors responsible for false ring formation in teak. Based on the anatomy and location within the true annual rings, four types of false rings were identified. The results from both cross-dating of plantation grown trees and controlled experiments, including induced drought, showed that rainfall during dry period, drought during active growing season, (container) polybag/field transplantation of seedlings and juvenility are the important causative factors of frequent false rings in teak. No definite relationship could be established between insect defoliation and the incidence of false rings. False ring formation in teak appears to be mainly a resultant feature of tree ring responses to different environmental and physical factors.     

Additive mixed models to study the effect of tree age and climatic factors on stem radial growth of Eucalyptus trees

The effect of tree age and climatic variables on stem radial growth of two hybrid clones of Eucalyptus was determined using longitudinal data from eastern South Africa.The stem radius of was measured weekly as the response variable.In addition to tree age,average weekly temperature,solar radiation,relative humidity and wind speed were simultaneously recorded with total rainfall at the site.An additive mixed effects model that incorporates a non-parametric smooth function was used.The results of the analysis indicate that the relationship between stem radius and each of the covariates can be explained by nonlinear functions.Models that account for the effect of clone and season together with their interaction in the parametric part of the additive mixed model were also fitted.The interaction between clone and season was not significant in all cases.For analyzing the joint effect all the covariates,additive mixed models that included two or more covariates were fitted.A significant effect of tree age was found in all cases.Although tree age was the key determinant of stem radial growth,weather variables also had a significant effect that was dependent on season.     

侧柏年轮宽度和年轮密度对气候变化的响应

通过响应函数分析和多元回归分析等年轮气候学方法,研究北京地区侧柏年轮宽度和年轮密度对气候变化的响应.结果表明:侧柏年轮宽度序列和年轮密度序列对气候变化相当敏感.前一年10月到当年9月的月平均气温和月降水量的响应函数模型可解释年轮宽度变化的58.45%,年轮密度变化的77.81%.夏季降水对侧柏年轮宽度有显著的正效应,而对年轮密度则表现出负效应.与降水的影响相反,夏季气温对侧柏年轮宽度表现出显著的负效应,对年轮密度表现出显著的正效应.其中,当年7月的气温和降水对侧柏年轮宽度和年轮密度的作用尤为显著.说明侧柏的年轮宽度和年轮密度均可用于年轮气候学研究,且年轮密度包含的信息量高于年轮宽度.     

Quantifying the effects of soil and climate on aboveground biomass production of <Emphasis Type="Italic">Salix miyabeana</Emphasis> SX67 in Quebec

Soil and climatic conditions for optimizing aboveground biomass yields of bioenergy short rotation coppices (SRCs) of Salix are not well elucidated. The objective of this study was to identify and quantify the limitations induced by soil and climate, and compare the magnitude of their effects, on annual aboveground yields across ten SRCs of Salix miyabeana SX67 in Quebec, Canada. The effects of weather variation between years on yields were also tested within locations. In five plots per SRC, soil bulk density, particle size, exchangeable cations and bulk composition were analysed, and moisture deficits were estimated using leaf δ¹³C. For each location, numerous weather variables were simulated for spring, summer and the whole growing season. Climate was calculated by averaging weather variables for growing seasons for which annual yields were available. Annual aboveground biomass yields were modelled using linear regression, partitioning of the variance and mixed models with soil, weather and climate variables as predictors. Across SRCs, silt content, soil organic matter, pH, exchangeable Ca and Mg, and total N and Zn were significantly and positively related to aboveground yields (adj. R ² ranging from 0.38 to 0.79). Generally, annual yields were negatively related to summer temperature within SRCs (adj. R ² = 0.92) and drought across SRCs (adj. R ² = 0.54). Partitioning of the variance revealed that soil variables (~80%) had a greater effect on productivity than did climate variables (~10%). In fact, soil properties buffered or exacerbated water shortages and, thus, had a preponderant effect on yield.     

Coast live oak, Quercus agrifolia, susceptibility and response to goldspotted oak borer, Agrilus auroguttatus, injury in southern California

Oak mortality is often associated with a complex of decline factors. We describe the morphological and physiological responses of coast live oak, Quercus agrifolia Née, in California to an invasive insect, the goldspotted oak borer (GSOB), Agrilus auroguttatus Schaeffer (Coleoptera: Buprestidae), and evaluate drought as a potential inciting factor. Morphological traits of 356 trees were assessed and physiological traits of 70 of these were monitored intensively over one growing season. Morphological characteristics of tree health included crown thinning and dieback; bole staining resulting from larval feeding; density of GSOB adult exit holes; and holes caused by woodpecker feeding. These characteristics were used to rank GSOB infestation/injury into four classes, and taken together, they explained 87% of the variation in a principal component analysis. Drought stress on various size/age and infestation classes of Q. agrifolia was measured by assessing branchlet pre-dawn and solar noon xylem water potential, leaf cell turgor potential, and water use efficiency over one growing season. Both morphological and physiological traits were highly variable in mature and old growth trees. Early summer plant water status (branchlet xylem water potential and water use efficiency) was similar between uninfested and newly colonized trees, suggesting that GSOB are not pre-selecting drought-stressed Q. agrifolia for oviposition. By late summer, leaf water and cell turgor potentials were lower in infested than in uninfested mature trees, suggesting that GSOB infestation causes drought stress in these trees. Among the tree size/age classes, infested old growth trees exhibited the greatest change in water use efficiency over the growing season, and showed greater morphological injury symptoms of decline than infested mature trees. Morphological attributes of decline in Q. agrifolia associated with GSOB were correlated weakly with increasing physiological drought stress among infestation classes of trees. We propose that the collection of morphological responses of Q. agrifolia to GSOB described here can be used to monitor the future expansion of the GSOB distribution as well as the GSOB-induced decline of Q. agrifolia in California.     

Electric resistance as a measure of tree water status during seasonal drought in a tropical dry forest in Costa Rica

Variation in electric resistance of stem tissues was used to measure differences and changes in water status among trees in a tropical dry forest in Costa Rica during the dry season. For more than 30 tree species, stem water content (SWC), measured as electric resistance between nails driven 20 mm deep into tree trunks, correlated well with wood density, saturation water content, dehydration, measured with the pressure chamber, and tree development during drought. At dry sites, SWC was lowest in hardwood trees (characterized by high wood density) and highest in stem-succulent lightwood trees (characterized by low wood density). Among hardwood trees, SWC varied with soil water availability. During the dry season, SWC declined before leaf shedding and increased during rehydration preceding bud break. The time course of seasonal changes in SWC apparently constitutes an indirect measure of variation in the relative water content of outer stem tissues, which determines development of dry-forest trees during the dry season.     

Sap flow index as an indicator of plant water status

Night and especially predawn tree water status is an important indicator of drought stress in trees. Leaf water potential (LWP) is frequently used as a measure of tree water status and hence drought stress; however, there are difficulties associated with sampling foliage from tall trees and determining LWP automatically. The current study was undertaken to determine whether sap flow index (SFI), which can be automatically and continuously recorded even during very low flows, can be used to estimate drought stress in trees caused by dry air under non-limiting soil water conditions. We made simultaneous measurements of LWP, heat pulse velocity (HPV) and SFI on apple trees (Malus domestica Borkh.) in the semiarid climate of southern Ukraine over several growing seasons. Predawn values of LWP were highly correlated with SFI. Over the range of low sap flow rates occurring at nighttime, where other methods of measuring sap flow are not sensitive, the SFI method was linear and very sensitive. Additional information about tree water status was obtained by comparing nighttime and daytime values of SFI. The ratio of predawn SFI to midday SFI and the period between the two daily SFI maxima (the first SFI peak occurred in the morning and the second peak occurred in the evening on cloudless days) can be used to characterize internal plant water balance. Although the daily course of SFI was variable, specific patterns were identified that reflected particular stages in the development of plant drought stress. An "air-drought-stress curve" was used to characterize the development of water stress in trees subjected to air drought during the growing season.     

Diversity of deciduousness and phenological traits of key Indian dry tropical forest trees

? In seasonally dry tropical forests deciduousness (leaflessness) is an important strategy of trees to survive in water stress period during summer. Deciduousness is a reflection of interacted effect of seasonal drought, tree characteristics and soil moisture conditions.

? The present study aims to document the diversity in leaf pheno-phases in terms of duration of deciduousness (which is reciprocal to growing season length), wood density, leaf mass per area (LMA) and leaf strategy index in 24 important tree species growing in the Vindhyan dry tropical forest in India.

? On the basis of phenological observations, the tree species were categorized into two main groups: leaf exchanging species exhibiting overlapping periods of leaf fall and leaf flush, and deciduous species whose timings of leaf flush and leaf fall differ resulting in a time lag (deciduousness) between the completion of leaf fall and initiation of leaf flush. Presence of wide range of deciduousness duration (from ca. a week to 7 months) among dry tropical trees indicates large variations in their growing season length. In the tree species studied, as the duration of deciduousness increased, leaf flushing period decreased significantly but leaf fall period showed little variation.

? Differing deciduousness in tree species exhibited substantial differences in their leafing (vegetative growth) pattern, as reflected by ratio of durations of leaf flush to leaf fall (leaf strategy index). Across different species, duration of deciduousness was significantly positively correlated with leaf strategy index, and significantly negatively correlated with both wood density and LMA.

? Wide variations in deciduousness, leaf strategy index, wood density and LMA in the 24 species investigated indicate considerable functional diversity in tree species growing in Vindhyan dry tropical region. Variation in seasonal duration of deciduousness among species is reflections of differences in tree functional traits like stem wood density, leaf strategy index and LMA.

     

大青山米老排树干径向生长变化及其对环境因子的响应

【目的】研究南亚热带大青山林区乡土珍贵树种米老排径向生长日变化特征及对环境因子的响应,为理解米老排生长-气候响应的生理机制、森林培育和保护等提供基础数据。【方法】利用点状树木径向生长变化记录仪对米老排径向生长变化进行连续观测,采用气象站同步测量辐射总量、土壤含水量、相对空气湿度、空气温度等环境因子数据。通过径向生长与环境因子的相关分析,研究米老排生长季第一个高峰期(5月)的不同分化等级径向生长日变化规律、不同天气条件下生长变化及径向生长-环境因子相互关系。【结果】1)不同分化等级米老排5月份的径向生长日变化量优势木(17.08μm)>平均木(13.94μm)>被压木(-1.81μm),差异极显著(P<0.01),且不同阶段的起始时间、阶段时长、变化量也有所不同。2)不同天气条件下,三种分化等级米老排径向日净生长量和日变化幅度均表现为雨天>晴天>阴天,受雨水影响,雨天条件下径向变化量远超其自身生长量;同种天气条件下,径向日净生长量优势木最大,日变化幅度则表现为平均木最大。3)不同分化等级米老排径向生长与相对空气湿度、降雨量呈极显著正相关关系,与气温、总辐射呈极显著负相关关系,而20 cm土壤含水率无显著性相关,且各影响因子均存在不同时间滞后效应。【结论】不同分化等级米老排由于林木生长状况的差异,导致其受到环境因子影响程度的不同,进而使得其径向生长变化量产生差异。     

Assessment of tree response to drought: validation of a methodology to identify and test proxies for monitoring past environmental changes in trees

A thinning experiment stand at Avoca, Ballinvalley, on the east coast of the Republic of Ireland was used to test a developed methodology aimed at monitoring drought stress, based on the analysis of growth rings obtained by coring. The stand incorporated six plots representing three thinning regimes (light, moderate and heavy) and was planted in the spring of 1943 on a brown earth soil. Radial growth (early- and latewood) was measured for the purpose of this study. A multidisciplinary approach was used to assess historic tree response to climate: specifically, the application of statistical tools such as principal component and canonical correlation analysis to dendrochronology, stable isotopes, ring density proxy, blue reflectance and forest biometrics. Results showed that radial growth was a good proxy for monitoring changes to moisture deficit, while maximum density and blue reflectance were appropriate for assessing changes in accumulated temperature for the growing season. Rainfall also influenced radial growth changes but not significantly, and was a major factor in stable carbon and oxygen discrimination, mostly in the latewood formation phase. Stable oxygen isotope analysis was more accurate than radial growth analysis in drought detection, as it helped detect drought signals in both early- and latewood while radial growth analysis only detected the drought signal in earlywood. Many studies have shown that tree rings provide vital information for marking past climatic events. This work provides a methodology to better identify and understand how commonly measured tree proxies relate to environmental parameters, and can best be used to characterize and pinpoint drought events (variously described using parameters such as like moisture deficit, accumulated temperature, rainfall and potential evaporation).     

The long way down--are carbon and oxygen isotope signals in the tree ring uncoupled from canopy physiological processes?

The carbon (δ(13)C) and oxygen (δ(18)O) stable isotope composition is widely used to obtain information on the linkages between environmental drivers and tree physiology over various time scales. The tree-ring archive can especially be exploited to reconstruct inter- and intra-annual variation of both climate and physiology. There is, however, a lack of information on the processes potentially affecting δ(13)C and δ(18)O on their way from assimilation in the leaf to the tree ring. As a consequence, the aim of this study was to trace the isotope signals in European beech (Fagus sylvatica L.) from leaf water (δ(18)O) and leaf assimilates (δ(13)C and δ(18)O) to tree-ring wood via phloem-transported compounds over a whole growing season. Phloem and leaf samples for δ(13)C and δ(18)O analyses as well as soil water, xylem water, leaf water and atmospheric water vapour samples for δ(18)O analysis were taken approximately every 2 weeks during the growing season of 2007. The δ(13)C and δ(18)O samples from the tree rings were dated intra-annually by monitoring the tree growth with dendrometers. δ(18)O in the phloem organic matter and tree-ring whole wood was not positively related to leaf water evaporative enrichment and δ(18)O of canopy organic matter pools. This finding implies a partial uncoupling of the tree-ring oxygen isotopic signal from canopy physiology. At the same time, internal carbon storage and remobilization physiology most likely prevented δ(13)C in tree-ring whole wood from being closely related to intra-annual variation in environmental drivers. Taking into account the post-photosynthetic isotope fractionation processes resulting in alterations of δ(13)C and δ(18)O not only in the tree ring but also in phloem carbohydrates, as well as the intra-annual timing of changes in the tree internal physiology, might help to better understand the meaning of the tree-ring isotope signal not only intra- but also inter-annually.     

Dissimilar stem and leaf hydraulic traits suggest varying drought tolerance among co-occurring Eucalyptus grandis × E. urophylla clones

In South Africa, there is currently a phenomenon where a Eucalyptus grandis × E. urophylla clone is dying on a large scale, whereas co-occurring clones in the same plantation area remain healthy. No primary biotic pathogen has been isolated from these dying trees. However, the plantation region has endured severe drought conditions. We hypothesised that the specific E. grandis × E. urophylla clone dieback is due in part to the affected trees having an elevated drought vulnerability. We tested this hypothesis retrospectively by examining how the dying (branch dieback started) as compared with two healthy and co-occurring E. grandis × E. urophylla clones vary in branch xylem anatomy (water transport) and leaf stable carbon isotopes (gas exchange) across two levels of mean annual precipitation in the planted landscape. There was significant intra-hybrid variation in the range of studied hydraulic traits. It appeared that the dying clone is maladapted to drier field conditions relative to the two healthy growing clones. Individuals of the drought-susceptible clone had increased hydraulic conductivity, having both wider vessel lumens and larger lumen fractions, and also did not regulate leaf-level transpiration in the drier landscape as efficiently as the two unaffected clones (lower δ¹³C ratios). In turn, one of the healthy growing clones had a completely different hydraulic strategy, having higher wood density, lower lumen fractions, and higher δ¹³C ratios that are widely considered as hydraulically resilient during water stress. The other clone appeared to also gain drought tolerance via higher leaf-level water-use efficiency, although drought tolerance was less clear regarding xylem anatomy. We highlight how variation in relative drought vulnerability can be highly significant even between clones within a single hybrid group. Knowledge of this variation in plant hydraulics between commercially planted and closely related Eucalyptus trees would complement clone deployment programmes in a target landscape.     

Effects of elevated carbon dioxide and drought on the growth and physiology of clonal Sitka spruce plants (Picea sitchensis (Bong.) Carr.)

Two-year-old Sitka spruce (Picea sitchensis (Bong.) Carr.) plants from four clones were grown in naturally lit growth chambers for 6 months at either ambient (350 ppm) or ambient + 250 ppm (600 ppm) CO(2) concentration. Plants were grown in large boxes filled with peat, in a system that allowed the roots of individual plants to be harvested easily at the end of the growing season. Half of the boxes were kept well watered and half were allowed to dry out slowly over the summer. Plants growing in elevated CO(2) showed a 6.9% increase in mean relative growth rate compared to controls in the drought treatment and a 9.8% increase compared to controls in the well-watered treatment, though there was considerable variation in response among the different clones and water treatments. Rates of net CO(2) assimilation were higher and stomatal conductances were lower in plants grown in elevated CO(2) than in ambient CO(2) in both the well-watered and drought treatments. Both of these factors contributed to the doubling of instantaneous water use efficiency. The partitioning of biomass to roots was unaffected by elevated CO(2), but the ratio of needle mass/stems + branches mass decreased. Together with reduced stomatal conductance, this probably caused the observed increases in xylem pressure potentials with elevated CO(2).     

A technique to identify annual growth rings in Eucalyptus grandis usingannual measurements of diameter at breast height and gamma raydensitometry

Many eucalypt species do not show distinct growth rings because cambial activity does not show a strong response to seasonal variation in climate. Eucalyptus grandis, one of the most important commercial hardwood species in South Africa, is one such example of a species that does not have well-defined growth rings. The light and dark bands visible on the cross-section of the wood of E. grandis do not always correspond with the growing season, which makes it difficult to resolve wood property data into annual increments. A method was developed to separate growth rings on wood density profiles of E. grandis into annual increments using bark-to-pith density profiles and annual measurements of diameter at breast height (DBH) from permanent sample plot (PSP) datasets. Using the PSP data, it was possible to assess the annual pattern of stem diameter growth at a compartment level by calculating the radial increment (RI) per year and expressing that value as a percentage of the radius at the end of the increment for that year. Mean radial increment percentage (%MRI) was calculated for each year and used to predict annual RI at an individual tree level. Predicted RI values for each tree were expressed as cumulative distances from the bark end and superimposed onto their respective density profiles. Predicted RI corresponded well with latewood density peaks and these separation points were considered a reliable guide to divide the density profile into annual increments closer to the bark end and into broader age classes closer to the pith. By assessing the pattern of variation in radial density within the context of the growth history of a compartment by means of annual PSP data, it was possible to confirm that growth rings on density profiles of E. grandis closer to the bark end can serve as a reliable representation of annual growth.