Does canopy mean nitrogen concentration explain variation in canopy light use efficiency across 14 contrasting forest sites?

The maximum light use efficiency (LUE?=?gross primary production (GPP)/absorbed photosynthetic photon flux density (aPPFD)) of plant canopies has been reported to vary spatially and some of this variation has previously been attributed to plant species differences. The canopy nitrogen concentration [N] can potentially explain some of this spatial variation. However, the current paradigm of the N-effect on photosynthesis is largely based on the relationship between photosynthetic capacity (A(max)) and [N], i.e., the effects of [N] on photosynthesis rates appear under high PPFD. A maximum LUE-[N] relationship, if it existed, would influence photosynthesis in the whole range of PPFD. We estimated maximum LUE for 14 eddy-covariance forest sites, examined its [N] dependency and investigated how the [N]-maximum LUE dependency could be incorporated into a GPP model. In the model, maximum LUE corresponds to LUE under optimal environmental conditions before light saturation takes place (the slope of GPP vs. PPFD under low PPFD). Maximum LUE was higher in deciduous/mixed than in coniferous sites, and correlated significantly with canopy mean [N]. Correlations between maximum LUE and canopy [N] existed regardless of daily PPFD, although we expected the correlation to disappear under low PPFD when LUE was also highest. Despite these correlations, including [N] in the model of GPP only marginally decreased the root mean squared error. Our results suggest that maximum LUE correlates linearly with canopy [N], but that a larger body of data is required before we can include this relationship into a GPP model. Gross primary production will therefore positively correlate with [N] already at low PPFD, and not only at high PPFD as is suggested by the prevailing paradigm of leaf-level A(max)-[N] relationships. This finding has consequences for modelling GPP driven by temporal changes or spatial variation in canopy [N].     

Leaf traits in relation to crown development, light interception and growth of elite families of loblolly and slash pine

Crown architecture and size influence leaf area distribution within tree crowns and have large effects on the light environment in forest canopies. The use of selected genotypes in combination with silvicultural treatments that optimize site conditions in forest plantations provide both a challenge and an opportunity to study the biological and environmental determinants of forest growth. We investigated tree growth, crown development and leaf traits of two elite families of loblolly pine (Pinus taeda L.) and one family of slash pine (P. elliottii Mill.) at canopy closure. Two contrasting silvicultural treatments -- repeated fertilization and control of competing vegetation (MI treatment), and a single fertilization and control of competing vegetation treatment (C treatment) -- were applied at two experimental sites in the West Gulf Coastal Plain in Texas and Louisiana. At a common tree size (diameter at breast height), loblolly pine trees had longer and wider crowns, and at the plot-level, intercepted a greater fraction of photosynthetic photon flux than slash pine trees. Leaf-level, light-saturated assimilation rates (A(max)) and both mass- and area-based leaf nitrogen (N) decreased, and specific leaf area (SLA) increased with increasing canopy depth. Leaf-trait gradients were steeper in crowns of loblolly pine trees than of slash pine trees for SLA and leaf N, but not for A(max). There were no species differences in A(max), except in mass-based photosynthesis in upper crowns, but the effect of silvicultural treatment on A(max) differed between sites. Across all crown positions, A(max) was correlated with leaf N, but the relationship differed between sites and treatments. Observed patterns of variation in leaf properties within crowns reflected acclimation to developing light gradients in stands with closing canopies. Tree growth was not directly related to A(max), but there was a strong correlation between tree growth and plot-level light interception in both species. Growth efficiency was unaffected by silvicultural treatment. Thus, when coupled with leaf area and light interception at the crown and canopy levels, A(max) provides insight into family and silvicultural effects on tree growth.     

Three-dimensional light structure of an upland Quercus stand post-tornado disturbance

Light is the most common limiting factor in forest plant communities,influencing species composition,stand structure,and stand productivity in closed canopy stands.Stand vertical light structure is relatively simple under a closed canopy because most light is captured by overstory trees.However,wind disturbance events create canopy openings from local to landscape scales that increase understory light intensity and vertical light structural complexity.We studied the effects of an EF-1 tornado on horizontal and vertical(i.e.three-dimensional)light structure within a Quercus stand to determine how light structure changed with increasing disturbance severity.We used a two-tiered method to collect photosynthetic photon flux density at 4.67 m and 1.37 m above the forest floor to construct three-dimensional light structure across a canopy disturbance severity gradient to see if light intensity varied with increasing tornado damage.Results indicate that increased canopy disturbance closer to the tornado track increased light penetration and light structure heterogeneity at lower forest strata.Increased light intensity correlated with increased sapling density that was more randomly distributed across the plot and had shifted light capture higher in the stand structure.Light penetration through the overstory was most strongly correlated with decreased stem density in the two most important tree species(based on relative dominance and relative density)in the stand,Quercus alba L.(r=-0.31)and Ostrya virginiana(Mill.)K.Koch(r=-0.27,p<.01),and indicated that understory light penetration was most affected by these two species.As managers are increasingly interested in patterning silvicultural entries on natural disturbances,they must understand residual stand and light structures that occur after natural disturbance events.By providing spatial light data that quantifies light structure post-disturbance,managers can use these results to improve planning required for long-term management.The study also provides comparisons with anthropogenic disturbances to the midstory that may offer useful comparisons to natural analogs for future silvicultural consideration.     

Effects of multiple factors on plant diversity of forest fragments in intensive farmland of Northern Portugal

The joint and independent effects of dominant tree species, forest patch spatial attributes, and forest structure and management as drivers of plant species richness and composition in small forest patches scattered within an intensive agricultural landscape were addressed.In a landscape with scattered urban and intensive dairy agricultural areas in north-west Portugal, within which small forest patches (dominated by pines, eucalypts, or both) represent semi-natural habitat islands, 50 small forest patches, with areas ranging between 0.3 and 3 ha, were selected and surveyed for vascular plant diversity, within dairy farming landscape mosaics dominated by annual forage crops. Explanatory variables were composed of three datasets derived either from GIS mapping or field observations: forest type (dominant tree species), forest patch spatial attributes (patch area and shape index), and measures of forest management and structure (diameter at breast height, tree density per hectare, and percentage cover of vegetation strata). Variations in these forest patch attributes were assessed across forest types, and related to measures of plant diversity (total, native, alien, woody, and herbaceous species richness). Redundancy analysis with variance partitioning was applied to evaluate the joint and independent effects of the three sets of variables on species assemblages. The recent shift in canopy dominance from pine to eucalypt observed in the region appears to be related to a (nonsignificant) tendency for the increase of patch area and to the decrease of patch complexity, as well as several changes in forest structure and management, expressed as a trend to denser tree canopies and lower cover of understory plants. Dominant tree species and attributes related to forest structure and management were the most important factors determining plant diversity. The joint effect of the dominant tree species and forest structure and management resulted in lower levels of plant species richness in eucalypt plantations. These were also more prone to invasion by alien species, probably due to decreased biotic resistance from unsaturated native plant assemblages. Our results draw attention to the importance of dominant tree species and management practices for the maintenance of plant diversity levels (species richness and composition) in dairy landscape mosaics, highlighting the importance of the remnants of semi-natural forests as refuges for plant diversity in the landscape context. Nonetheless, forest plant diversity could further be fostered by promoting naturalness of pine stands and the regeneration of native oak woodlands in some forest areas. This would also diversify the range of ecosystem services that could be provided by forest areas in these peri-urban farmlands.     

Photosynthesis and respiration of black spruce at three organizational scales: shoot, branch and canopy

To gain insight into the function of photosynthesis and respiration as processes operating within a global ecosystem, we measured gas exchange of mature black spruce (Picea mariana (Mill.) B.S.P.) trees at three organizational scales: individual shoots, whole branches and a forest canopy. A biochemical model was fitted to these data, and physiological parameters were extracted. Pronounced seasonal variation in the estimated model parameters was found at all three organizational scales, highlighting the need to make physiological measurements throughout the year. For example, it took over 100 days for physiological activity to increase from zero during the springtime thaw to its yearly maximum. Good agreement was found between parameter values estimated for the different organizational scales, suggesting that, in the case of aerodynamically rough, largely mono-specific forest canopies, physiological parameters can be estimated from eddy covariance flux measurements. The small differences between photosynthetic parameters estimated at the different scales suggest that the overall spatial organization of photosynthetic capacity is nearly optimized for carbon uptake at each scale.     

林隙微生境及更新研究进展

系统评述了国内外关于林隙干扰及更新的最新研究进展.林隙是一种经常发生的小规模干扰,是森林群落时空格局变化的驱动力.不同的林隙干扰状况导致了林隙内光照的差异,进一步造成林隙内温度、湿度、土壤理化性质的变化,形成了微生境的时空异质性.林隙干扰通过改变微生境而对幼苗的出土、存活、定居和生长产生重要影响.林隙内种子库的物种丰富度变化很大,是幼苗分布格局的重要影响因子.更新幼苗的存活率和生长量受林隙大小及幼苗在林隙中位置的影响.指出了这一领域今后的研究方向.     

Decreasing the error in the measurement of the ecosystem effective leaf area index of a Pinus massoniana forest

Decreasing the forest ecosystem leaf-area index error(LAIe)helps accurately estimate the growth and light energy utilization of aboveground foliage.Analyzing light transmission in forest ecosystems can effectively determine LAIe.The LAI-2200 plant canopy analyzer(PCA)is used extensively for rapid field-effective LAI(LAIe)measurements and primarily to measure forest canopy LAIe values.However,sometimes this parameter must also be measured in forests with small clearings.In this study,we used the LAI-2200 PCA to obtain one A-value and four B-values each for the canopy,herbaceous layer,and forest ecosystem LAIe.Field measurements showed that the three LAIe types were obviously different.In certain quadrats,the average herbaceous layer(Dicranopteris dichotoma Bernh.)LAIe apparently exceeded that of the Pinus massoniana forest ecosystem.The sources of this error were measuring and recording A-value readings for small canopies and underestimating the ecosystem LAIe.We obtained similar coefficients of determination for both the pre-recomputation and post-recomputation of the canopy and forest ecosystem LAIe(R^2C 0.96 and R^2C 0.99,respectively);thus,the error was decreased.Measuring field LAIe with the LAI-2200 PCA and recomputation should compensate for LAIe underestimation in complex forest ecosystems.     

Spatial and temporal variations in soil respiration in relation to stand structure and soil parameters in an unmanaged beech forest

Soil CO2 efflux (soil respiration) plays a crucial role in the global carbon cycle and efflux rates may be strongly altered by climate change. We investigated the spatial patterns of soil respiration rates in 144 measurement locations in a 0.5-ha plot and the temporal patterns along a 300-m transect in the 0.5-ha plot. Measurements were made in an unmanaged, highly heterogeneous beech forest during 2000 and 2001. We investigated the effects of soil, roots and forest stand structure on soil respiration, and we also assessed the stability of these spatial patterns over time. Soil temperature alone explained between 68 and 95% of the temporal variation in soil respiration; however, pronounced spatial scatter of respiration rates was not explained by soil temperature. The observed spatial patterns stayed remarkably stable throughout the growing season and over 2 years. The most important structural parameter of the stand was the mean diameter at breast height of trees within a distance of 4 m of the measurement locations (m-dbh4), which explained 10-19% of the variation in soil respiration throughout the growing season. Among the soil chemical parameters, carbon content (bulk as well as dissolved) and magnesium content explained 62% of the spatial variation in soil respiration. The final best model combining soil, root and stand structural parameters (fine root biomass, soil carbon content, m-dbh4 and soil water content) explained 79% of the variation in soil respiration, illustrating the importance of both biotic and abiotic factors.     

Asymmetric competition increases leaf inclination effect on light absorption in mixed canopies

? Context

The effects of leaf inclination on plant light capture, growth, and water balance of monospecific canopies are well documented, but we still lack information on such effects in the case of multispecific canopies.

? Aims

We investigated the effects of leaf inclination on the absorption of photosynthetically active radiation (PAR) of a mixed forest.

? Methods

We ran a 3D mechanistic radiation transfer model for a Mediterranean forest where Pinus halepensis makes the upper strata while Quercus ilex occupies the lower strata. As factors, we included (1) the distributions of leaf inclinations that ranged from vertical to horizontal (including the actual inclinations), (2) the fraction of diffuse light, sun position, and leaf area index (LAI), and (3) the Pinus/Quercus LAI ratio.

? Results

Simulated PAR absorption was more than twice as sensitive to leaf inclination in oaks than in pines because oaks depended on PAR transmitted below the pine layer. The extent of the effect depended on season, fraction of diffuse light, LAI, and vegetation spatial structure. None of the observed inclinations maximized PAR absorption, suggesting a trade-off with water economy.

? Conclusion

Erroneous assumptions about leaf inclination lead to larger errors when modelling heterogeneous, mixed canopies. This also highlights potential caveats when using models that do not account for the spatial structure of canopies.     

Co-optimal distribution of leaf nitrogen and hydraulic conductance in plant canopies

Leaf properties vary significantly within plant canopies, due to the strong gradient in light availability through the canopy, and the need for plants to use resources efficiently. At high light, photosynthesis is maximized when leaves have a high nitrogen content and water supply, whereas at low light leaves have a lower requirement for both nitrogen and water. Studies of the distribution of leaf nitrogen (N) within canopies have shown that, if water supply is ignored, the optimal distribution is that where N is proportional to light, but that the gradient of N in real canopies is shallower than the optimal distribution. We extend this work by considering the optimal co-allocation of nitrogen and water supply within plant canopies. We developed a simple 'toy' two-leaf canopy model and optimized the distribution of N and hydraulic conductance (K) between the two leaves. We asked whether hydraulic constraints to water supply can explain shallow N gradients in canopies. We found that the optimal N distribution within plant canopies is proportional to the light distribution only if hydraulic conductance, K, is also optimally distributed. The optimal distribution of K is that where K and N are both proportional to incident light, such that optimal K is highest to the upper canopy. If the plant is constrained in its ability to construct higher K to sun-exposed leaves, the optimal N distribution does not follow the gradient in light within canopies, but instead follows a shallower gradient. We therefore hypothesize that measured deviations from the predicted optimal distribution of N could be explained by constraints on the distribution of K within canopies. Further empirical research is required on the extent to which plants can construct optimal K distributions, and whether shallow within-canopy N distributions can be explained by sub-optimal K distributions.     

Comparative analysis of photosynthetic light environments within the crowns of juvenile rain forest trees

Irradiances within the crowns of saplings of two tropical tree species were simultaneously compared in primary rain forest in Costa Rica. The species examined, Minquartia guianensis Aubl., a relatively slow-growing, canopy species, and Pithecellobium pedicellare (DC) Benth., a less-tolerant, emergent species, have different crown and leaf display patterns. Crown light environments were assessed by placing arrays of quantum sensors among leaves and recording at 5-s intervals for seven days with microloggers. Median total daily quantum flux densities for saplings of both species were less than 2% of full sun and did not differ significantly. More than 90% of the measurements within the crowns of these saplings were less than 25 micromol m(-2) s(-1). Spatial variability of photon flux densities within sapling crowns was similar for the two species despite differences in leaf display patterns. In saplings of both species, photon flux densities varied significantly over the relatively short distances within crowns and from day to day. Height growth of both species was significantly correlated with total daily photon flux densities and with percentage of full sun. However, only the tolerant species, Minquartia, showed a significant correlation between diameter growth and crown light environment.     

Impact of the bambooPhyllostachys bambusoides on the light environment and plant communities on riverbanks

The bambooPhyllostachys bambusoides has been widely planted on riverbanks in Japan to protect against erosion. Recently, unmanagedP. bambusoides stands with high culm density are spreading along the banks of rivers. In order to study the impact of the bamboo stands on riparian ecosystems, the relations between bamboo stand structure, light conditions and plant communities under the canopy were examined. The study sites were set in bamboo stands with various culm densities in the middle of the Yahagi River in Toyota, Aichi Prefecture. There was a close relationship between canopy coverage and relative photon flux density (RPFD) in the bamboo stands. RPFD showed a highly negative relationship with total culm density, whereas no significant relation was observed between the basal area of bamboo and RPFD. The total number of plant species in the bamboo stands increased with decreasing culm density. Only a small number of tree saplings were observed in the stands with the highest culm densities, while several species of forest floor herbs were found in the stands with lower culm densities. A number of invasive and annual plants colonized the thinned stand. We suggest that moderate thinning and clearing of dead culms are needed to maintain biological diversity in the bamboo stands on riverbanks.     

A COMPARISON OF RAINFALL IN DIFFERENT WOODLANDS

At Bedgebury the rainfall in thirteen forest plots planted withdifferent tree species has been compared with that in the open.In all these forest plots the trees have closed canopy within20 years of planting and the canopies differ considerably intheir structure. From 6 to 93 per cent. of the gross rainfallmay be retained on the tree canopies and lost to the soil. Theproportion of precipitation intercepted is greatest in lightrains and least in heavy rains. During light showers considerablymore moisture is retained on the conifer than on the hardwoodcanopies. In the heavier showers rainwater flows down the stems,but stem flow represents a small proportion of the total waterreaching the forest floor. The amount of water which reachesthe ground along the trunks varies with the species and thecharacter of branching. Some raindrops penetrate directly throughthe canopy, but many are intercepted by the canopies and coalesceto fall as large drops on the forest floor. These large waterdropstend to fall continuously on the same spot so that in the plantationsthe water is distributed unevenly. In the winter months, whenthe deciduous species have shed their leaves, snow penetratesmore easily through their canopies than those of the evergreenconifers.     

Sunflecks in trees and forests: from photosynthetic physiology to global change biology

Sunflecks are brief, intermittent periods of high photon flux density (PFD) that can significantly improve carbon gain in shaded forest understories and lower canopies of trees. In this review, we discuss the physiological basis of leaf-level responses to sunflecks and the mechanisms plants use to tolerate sudden changes in PFD and leaf temperature induced by sunflecks. We also examine the potential effects of climate change stresses (including elevated temperatures, rising CO(2) concentrations and drought) on the ability of tree species to use sunflecks, and advocate more research to improve our predictions of seedling and tree carbon gain in future climates. Lastly, while we have the ability to model realistic responses of photosynthesis to fluctuating PFD, dynamic responses of photosynthesis to sunflecks are not accounted for in current models of canopy carbon uptake, which can lead to substantial overestimates of forest carbon fixation. Since sunflecks are a critical component of seasonal carbon gain for shaded leaves, sunfleck regimes and physiological responses to sunflecks should be incorporated into models to more accurately capture forest carbon dynamics.     

Spatial variation of local stand structure in an Abies forest, 45 years after a large disturbance by the Isewan typhoon

Heterogeneity of forest stand structure often results from repeated small disturbances, but structural variation also arises in a stand that has regenerated after a single large stand-replacing disturbance. We explored the structural variation within a subalpine Abies forest in Japan that regenerated after a large typhoon in 1959. In 2004, four 50 × 50 m plots were established at two sites in the regenerated forest. To characterize local stand structure within each plot, we determined the stem density, stand basal area, mean diameter at breast height (DBH), and coefficient of variation (CV) of DBH in 10 × 10 m subplots. We analyzed the spatial distribution pattern of the trees in each subplot using the L(t) function and categorized the distributions as clumped or non-clumped. The analysis revealed marked variation among subplots in the stand’s structural characteristics. Although the spatial distribution patterns of the trees in all plots were clumped, 11 of 50 subplots at one site and 39 of 50 subplots at the other site were non-clumped. Subplots with a clumped distribution pattern generally had a smaller basal area, smaller mean DBH, and greater CV of DBH than subplots with the same density but a non-clumped pattern. These results illustrated the spatial heterogeneity of forest structure that can arise in Abies forest that has experienced a large disturbance, probably because of the different densities and distribution of saplings surviving the disturbance and the different forest growth dynamics that result.     

Oak restoration in water-limited pine plantations: interactive effects of overstory light interception and water availability on understory oak performance

Oak regeneration within pine monocultures is an opportunity to diversify forest structure. We examined the relationships between overstory (Pinus brutia) light interception and understory oak (Quercus ithaburensis) performance in water-limited forests. The study was performed in a mature pine plantation in Mediterranean Israel. Twenty-year-old oaks differing in location with respect to pine overstory and representing a gradient of light availability, such as open space (irradiance 100 %), interface (17–77 %), and understory (14–23 %), were monitored. Photosynthetic photon flux density (PPFD), leaf gas exchange, and twig water potential (TWP) were measured during the growth season under increasing drought stress. Predawn TWP decreased sharply from early to late spring and was positively related to irradiance during mid-spring only. Predawn to midday TWP gradient was positively related to irradiance mostly so during mid-spring. Daily averages of stomatal conductance (gs), net carbon assimilation rate (A), and transpiration rate (E) were highest in early spring and decreased gradually toward late spring. They were positively related to irradiance though this relationship became less pronounced from early to late spring. Oak height and stem basal area were positively related to irradiance. A/gs ratio was positively related to irradiance throughout the entire growth season. It increased from early to mid-spring but decreased toward late spring. A/PPFD ratio decreased from early to late spring showing a negative relationship with irradiance. We concluded that light availability was mainly responsible for spatial variation in oak performance and proposed that small-scale overstory gaps aiming for direct sunlight exposure during early spring should achieve maximum understory oak performance with minimal pine removal.     

Short-term light and leaf photosynthetic dynamics affect estimates of daily understory photosynthesis in four tree species

Instantaneous measurements of photosynthesis are often implicitly or explicitly scaled to longer time frames to provide an understanding of plant performance in a given environment. For plants growing in a forest understory, results from photosynthetic light response curves in conjunction with diurnal light data are frequently extrapolated to daily photosynthesis (A(day)), ignoring dynamic photosynthetic responses to light. In this study, we evaluated the importance of two factors on A(day) estimates: dynamic physiological responses to photosynthetic photon flux density (PPFD); and time-resolution of the PPFD data used for modeling. We used a dynamic photosynthesis model to investigate how these factors interact with species-specific photosynthetic traits, forest type, and sky conditions to affect the accuracy of A(day) predictions. Increasing time-averaging of PPFD significantly increased the relative overestimation of A(day) similarly for all study species because of the nonlinear response of photosynthesis to PPFD (15% with 5-min PPFD means). Depending on the light environment characteristics and species-specific dynamic responses to PPFD, understory tree A(day) can be overestimated by 6-42% for the study species by ignoring these dynamics. Although these overestimates decrease under cloudy conditions where direct sunlight and consequently understory sunfleck radiation is reduced, they are still significant. Within a species, overestimation of A(day) as a result of ignoring dynamic responses was highly dependent on daily sunfleck PPFD and the frequency and irradiance of sunflecks. Overall, large overestimates of A(day) in understory trees may cause misleading inferences concerning species growth and competition in forest understories with < 2% full sunlight. We conclude that comparisons of A(day) among co-occurring understory species in deep shade will be enhanced by consideration of sunflecks by using high-resolution PPFD data and understanding the physiological responses to sunfleck variation.     

Temperature, light and leaf hydraulic conductance of little-leaf linden (Tilia cordata) in a mixed forest canopy

Response of whole-leaf hydraulic conductance (G(L)) in little-leaf linden (Tilia cordata Mill.) to temperature and photosynthetic photon flux (Q(P)) was estimated by the evaporative flux method under natural conditions in a mixed forest canopy. Mean midday G(L) in the lower- and upper-crown foliage was 1.14 and 3.06 mmol m(-2) s(-1) MPa(-1), respectively. Over the study period, leaf temperature (T(L)) explained about 67% of the variation in G(L), and Q(P) explained about 10%. Leaf water potential and crown position also affected G(L) significantly. About a third of the temperature effect was attributable to changes in the viscosity of water, and two thirds to changes in protoplast permeability (i.e., symplastic conductance). Leaf hydraulic conductance was highly sensitive to changes in Q(P) when Q(P) was less than 200 micromol m(-2) s(-1), and G(L) sensitivity decreased with increasing irradiance. Sensitivity of G(L) to variation in T(L) increased consistently with increasing temperature in the range of 16 to 29 degrees C. There were positive interactions between temperature and light in their effects on G(L): the light response was more pronounced at higher leaf temperatures. Because of frequent rains during the study period, the trees experienced no soil water deficit, and, within the range experienced, soil water potential had no effect on G(L). Leaf hydraulic conductance exhibited a seasonal pattern that could be explained primarily by temporal variability in mean air temperature and irradiance, in addition to which an age-related trend (P<0.001) of increasing G(L) from the end of June to the beginning of August was observed.     

Effects of shade on growth, biomass allocation and leaf morphology in European yew (Taxus baccata L.)

The impact of shade on the growth of European yew (Taxus baccata L.) saplings was investigated over a three-year period using artificial shading to simulate four different light regimes (3, 7, 27 and 100 % relative photosynthetic photon flux density, RPPFD). There was no mortality attributable to shading even under the 3 % RPPFD treatment. Increasing shade was positively associated with specific leaf area, leaf length, leaf width and total chlorophyll content, but negatively associated with plant height, stem diameter, total dry weight and root to leaf and shoot ratio. Discoloration of the foliage occurred in plants grown in 100 % RPPFD conditions (resulting in reduced growth rates) and those transferred to 100 % RPPFD conditions after being shade-acclimated for 2 years. Evidence suggests that T. baccata has the ability to regenerate beneath a lighter canopy but beneath denser canopies gap dynamics will play an important role in facilitating successful regeneration and this needs to be reflected in management of natural populations of this declining species.