Photosynthetic capacity in relation to nitrogen in the canopy of a Quercus robur, Fraxinus angustifolia and Tilia cordata flood plain forest

We measured gas exchange and various leaf parameters of ash (Fraxinus angustifolia Vahl.) and oak (Quercus robur L.) in the high canopy and of lime (Tilia cordata Mill.) in the lower canopy of a planted, 120-year-old floodplain forest in southern Moravia, Czech Republic. The high-canopy leaves of F. angustifolia and Q. robur had nitrogen concentrations on a leaf area basis (N(area)) that were twice those of low-canopy leaves of T. cordata. Upper-canopy leaves of F. angustifolia had a photosynthetic rate at light saturation (A(max)) of about 16 micromol CO2 m(-2) s(-1), whereas A(max) of the upper-canopy foliage of Q. robur achieved only about two thirds of this value. Contrary to previous investigations of photosynthetic performance in monospecific stands, leaves of the uppermost branches of T. cordata at 15-m height had the highest A(max) and transpiration rate among the species studied. Water-use efficiency (WUE) was low in T. cordata at 15-m canopy height, whereas WUE was significantly higher for Q. robur leaves at 27-m height than for the other species. Leaves of T. cordata at 15-m height showed the strongest relationship between A(max) and N(area) (R2 = 0.90) followed by F. angustifolia (R2 = 0.69). The strong correlation between photosynthesis and nitrogen concentration in T. cordata at 15 m, together with the steep regression slope for the A(max):N(area) relationship, indicated that nitrogen allocation to the photosynthetic apparatus resulted in high nitrogen-use efficiency of light-saturated photosynthesis (PNUE). Despite differences in PNUE among species, PNUE was fairly constant for leaves sampled from the same canopy position, suggesting that single-leaf parameters are matched to optimize PNUE for prevailing light conditions. High PNUE in T. cordata at 15 m partially compensated for the species' subordinate position in the canopy, and may be an important mechanism for its coexistence in highly structured vegetation.     

Effects of enhanced hydraulic supply for foliage on stomatal responses in little-leaf linden (<Emphasis Type="Italic">Tilia cordata</Emphasis> Mill.)

Responses of leaf conductance (g _L) to variation in environmental and plant hydraulic factors were examined on intact and detached shoots of little-leaf linden (Tilia cordata Mill.) with respect to branch position in the crown. Using detached shoots, we manipulated leaf water supply and light availability in order to separate the effects of insufficient hydraulic supply and low irradiance. The intact upper-crown leaves demonstrated 2.0–2.2 times higher (P < 0.001) daily maxima of g _L compared to the lower-crown leaves growing in the shadow of upper branches. Mean soil-to-leaf conductance (G _T) was 1.9 times higher (P < 0.001) for the upper-crown foliage compared to that of the lower crown. The total hydraulic resistance was distributed: soil to distal shoots—41–51%, 25-cm distal shoots—10–15% and leaves—39–44%. In lower branches, g _L was constrained by both low light availability and limited water supply; in upper branches, only by irradiance. Artificial reduction of hydraulic constraints raised bulk leaf water potential (Ψ _L) and made g _L less sensitive to changes in both atmospheric and plant factors. Stomatal responses to leaf-to-air vapour pressure difference (VPD) were significantly modified by leaf water status: high Ψ _L seemingly inverted the g _L versus VPD relationship. Enhanced water supply increased g _L and transpiration rate (E) in the lower-crown foliage, but not in the upper-crown foliage. The results support the idea that leaves in the lower canopy are hydraulically more constrained than those in the upper canopy.     

Responses of foliar photosynthetic electron transport, pigment stoichiometry, and stomatal conductance to interacting environmental factors in a mixed species forest canopy

We studied limitations caused by variations in leaf temperature and soil water availability on photosynthetic electron transport rates calculated from foliar chlorophyll fluorescence analysis (U) in a natural deciduous forest canopy composed of shade-intolerant Populus tremula L. and shade-tolerant Tilia cordata Mill. In both species, there was a positive linear relationship between light-saturated U (Umax) per unit leaf area and mean seasonal integrated daily quantum flux density (Ss, mol per square m per day). Acclimation of leaf dry mass per area and nitrogen per area to growth irradiance largely accounted for this positive scaling. However, the slopes of the Umax versus Ss relationships were greater on days when leaf temperature was high than on days when leaf temperature was low. Overall, Umax varied 2.5-fold across a temperature range of 20-30 degrees C. Maximum stomatal conductance (Gmax) also scaled positively with Ss. Although Gmax observed during daily time courses, and stomatal conductances during Umax measurements declined in response to seasonally decreasing soil water contents, was insensitive to prolonged water stress, and was not strongly correlated with stomatal conductances during its estimation. These results suggest that photorespiration was an important electron sink when intercellular CO2 concentration was low because of closed stomata. Given that xanthophyll cycle pool size (VAZ, sum of violaxanthin, antheraxanthin, and zeaxanthin) may play an important role in dissipation of excess excitation energy, the response of VAZ to fluctuating light and temperature provided another possible explanation for the stable Umax. Xanthophyll cycle carotenoids per total leaf chlorophyll (VAZ/Chl) scaled positively with integrated light and negatively with daily minimum air temperature, whereas the correlation between VAZ/Chl and irradiance was best with integrated light averaged over 3 days preceding foliar sampling. We conclude that the potential capacity for electron transport is determined by long-term acclimation of U to certain canopy light conditions, and that the rapid adjustment of the capacity for excitation energy dissipation plays a significant part in the stabilization of this potential capacity. Sustained high capacity of photosynthetic electron transport during stress periods provides an explanation for the instantaneous response of U to short-term weather fluctuations, but also indicates that U restricts potential carbon gain under conditions of water limitation less than does stomatal conductance.     

Use of thermal imaging to determine leaf conductance along a canopy gradient in European beech (Fagus sylvatica)

Using an infrared camera, we measured the leaf temperature across different canopy positions of a 23-m-tall deciduous forest tree (Fagus sylvatica L.) including typical sun and shade leaves as well as intermediate leaf forms, which differed significantly in specific leaf area (SLA). We calculated a temperature index (I(G)) and a crop water stress index (CWSI) using the surface temperatures of wet and dry reference leaves. Additional indices were computed using air temperature plus 5 °C (I(G)?+?5, CWSI?+?5) as dry references. The minimum temperature of the wet leaf and the maximum temperature of the dry leaf proved to be most suitable as reference values. We correlated the temperature indices with leaf area-related conductance to water vapor (g(L)) using porometry at the leaf level and using xylem sap flow at the branch level. At the leaf and at the branch level, I(G) and CWSI were equally well suited as proxies of g(L), whereas the relationships of I(G)?+?5 and CWSI?+?5 with g(L) were only weak or even insignificant. At the leaf level, the correlations of I(G) and CWSI with g(L) were significant in all parts of the crown. The slopes of g(L) vs. I(G) and CWSI did not differ significantly among the crown parts; this indicates that they were not influenced by SLA or irradiance. At the branch level, close correlations (r?>?0.8) were found between temperature indices and g(L) across the crown. These results demonstrate that satisfactory relationships between temperature indices and g(L) can be established in tall trees even in those canopy parts that are exposed to relatively low levels of irradiance and exhibit relatively low values of g(L).     

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.     

Diurnal patterns of leaf photosynthesis, conductance and water potential at the top of a lowland rain forest canopy in Cameroon: measurements from the Radeau des Cimes

Diurnal patterns of leaf conductance, net photosynthesis and water potential of five tree species were measured at the top of the canopy in a tropical lowland rain forest in southwestern Cameroon. Access to the 40 m canopy was by a large canopy-supported raft, the Radeau des Cimes. The measurements were made under ambient conditions, but the raft altered the local energy balance at times, resulting in elevated leaf temperatures. Leaf water potential was equal to or greater than the gravitational potential at 40 m in the early morning, falling to values as low as -3.0 MPa near midday. Net photosynthesis and conductance were typically highest during midmorning, with values of about 10-12 micro mol CO(2) m(-2) s(-1) and 0.2-0.3 mol H(2)O m(-2) s(-1), respectively. Leaf conductance and net photosynthesis commonly declined through midday with occasional recovery late in the day. Photosynthesis was negatively related to leaf temperature above midday air temperature maxima. These patterns were similar to those observed in other seasonally droughted evergreen communities, such as Mediterranean-climate shrubs, and indicate that environmental factors may cause stomatal closure and limit photosynthesis in tropical rain forests during the midday period.     

Diurnal and seasonal variations in leaf hydraulic conductance in evergreen and deciduous trees

We studied changes in the hydraulic conductance of leaves (K(leaf)) between dawn and dusk during the growth period (July) and at midday at the beginning of autumn in four tree species. The main objectives of the study were to check the extent of diurnal and seasonal changes in K(leaf) and the relationships between K(leaf), irradiance and leaf gas exchange. Two evergreen (Aleurites moluccana and Persea americana) and two deciduous trees (Platanus orientalis and Quercus rubra) were studied. Leaf hydraulic conductance was measured every 2 h between 0700 and 1900 h in July and compared with values measured between 0900 and 1300 h in October. Other variables measured were photosynthetically active radiation (PAR), leaf conductance to water vapor (gL) and water potential (psiL). In July, K(leaf) varied by up to 75% in Pe. americana on a diurnal basis and by at least 44% in Q. rubra. The diurnal time course of K(leaf) showed a distinct increase between dawn and late morning (1100 h) and a subsequent decrease in the evening in A. moluccana and Pl. orientalis, whereas in the other two species, K(leaf) was highest just after dawn and lowest in the evening. In October, K(leaf) of all the species studied was lower than in July, with differences of 20 to 28% for A. moluccana and Pl. orientalis and of 66 to over 70% in Pe. americana and Q. rubra, respectively. Significant correlations were found between PAR and K(leaf) (in all species) as well as between gL and K(leaf) (in three out of four species). Leaf habit (evergreen or deciduous) did not influence absolute values of K(leaf) or its diurnal variation.     

Gas exchange characteristics of a Canarian laurel forest tree species (Laurus azorica) in relation to environmental conditions and leaf canopy position

Diurnal courses of gas exchange were measured over a 1-year period in fully expanded current-year leaves in the upper (sun-exposed, 18 m above ground) and the lower (shaded, 12 m above ground) canopy of Laurus azorica (Seub.) Franco, a major canopy species of the Canarian laurel forest in Tenerife, Canary Islands, Spain. Laurus azorica exhibited high leaf plasticity in gas exchange characteristics, with a maximum carbon assimilation rate (Amax) of shade leaves about 50% that of sun leaves. This difference reflects the high leaf area index (LAI) of the stand and the correspondingly sharp light attenuation with increasing canopy depth. In sun leaves, Amax peaked at about 11 micromol m-2 s-1 and maximum transpiration (E) was about 8 mmol m-2 s-1, which corresponded with a maximum stomatal conductance (gs) of about 650 mmol m-2 s-1. Mean maximum instantaneous water-use efficiency (WUE) was 1.5 mmol mol-1 and the mean maximum A/gs was 20-35 micromol mol-1. Mean minimum internal CO2 concentration (Ci) was 225 micromol mol-1. Although high air vapor pressure deficit (VPD) caused a small decrease in gs, it remained high enough to maintain relatively high A and E. These gas exchange characteristics indicate a non-conservative use of water, which is appropriate for a species subject to droughts that are mild or of short duration. In this respect, Laurus azorica differs from its congener, L. nobilis L., of the Mediterranean region and other shrubs growing in Mediterranean-type climates in California and Chile that have to withstand more severe or more prolonged droughts.     

Variation in canopy structure, light and soil nutrition across elevation of a Sri Lankan tropical rain forest

Mixed dipterocarp forests are perhaps the single most important rain forest type in the wet tropics. Only a few studies have purposefully examined differences in resource availability across mixed dipterocarp forest landscapes by simply measuring the abiotic variables of light, soil nutrition and soil water availability in relation to forest structure. We sought to directly measure the environment of canopy gaps across elevation and geology—from lowland mixed dipterocarp forest (100 m amsl) to lower montane dipterocarp forest (1200 m amsl) in southwest Sri Lanka. Middle elevation gap sites (300–900 m amsl) were subdivided into valley, mid-slope and ridge topographic positions. Eighteen natural disturbances all of which were canopy openings caused by tree fall, were randomly selected within primary rain forest that ranged across 100–1200 m elevation. Plots were placed in gap centers and in adjacent understories and measurements taken of forest structure (basal area, canopy height, canopy cover index, CCI), shade (light sensors—photosynthetically active radiation [PAR], canopy hemispherical photographs—global site factor [GSF]) and soil nutrition (pH, exchangeable Al, K, Mg and Ca; Total N; and plant available P). Soil moisture was measured at bi-weekly intervals for five years across middle elevation sites only (300–900 m amsl). Stand basal area, mean canopy height, and canopy cover index all declined with increase in elevation. Understory PAR and GSF decreased with increases in canopy height, basal area and CCI. Size of canopy opening decreased with increase in elevation, but PAR and GSF increased. Valley sites had significantly greater levels of mean percent soil water content as compared to mid-slope and ridge sites of middle elevation sites. However, at the onset of the southwest monsoons in May all sites were similar. Differences were most pronounced during the dry season (December–April). No differences in soil moisture content could be found between gap and understory microsites. K and Ca in gap centers and adjacent forest understories increased with increase in elevation and change in associated geology. pH increased and Al decreased with elevation and associated geology but only for forest understory conditions. Results demonstrate strong differentiation in soil and light resources with elevation that appears related to size of tree-fall disturbance, stature of the forest, topographic position and underlying geology and soil-weathering environment. This suggests that forest management and conservation practices need to develop and tailor techniques and treatments (silviculture) to the forest that emulate and/or account for change in elevation, geology and topographic position. Further studies are needed to identify which are the primary underlying mechanisms (e.g. temperature, wind, soil nutrients, soil moisture availability) defining change in forest structure across elevation.     

Interspecific variation in nighttime transpiration and stomatal conductance in a mixed New England deciduous forest

Transpiration is generally assumed to be insignificant at night when stomata close in response to the lack of photosynthetically active radiation. However, there is increasing evidence that the stomata of some species remain open at night, which would allow for nighttime transpiration if there were a sufficient environmental driving force. We examined nighttime water use in co-occurring species in a mixed deciduous stand at Harvard Forest, MA, using whole-tree and leaf-level measurements. Diurnal whole-tree water use was monitored continuously with Granier-style sap flux sensors in paper birch (Betula papyrifera Marsh.), red oak (Quercus rubra L.) and red maple (Acer rubrum L.). An analysis was conducted in which nighttime water flux could be partitioned between refilling of internal water stores and transpiration. Substantial nighttime sap flux was observed in all species and much of this flux was attributed to the refilling of depleted water stores. However, in paper birch, nighttime sap flux frequently exceeded recharge estimates. Over 10% of the total daily sap flux during the growing season was due to transpiration at night in paper birch. Nighttime sap flux was over 8% of the total daily flux in red oak and 2% in red maple; however, this flux was mainly associated with recharge. On nights with elevated vapor pressure deficit, sap flux continued through the night in paper birch, whereas it reached zero during the night in red oak and red maple. Measurements of leaf-level gas exchange on a night with elevated vapor pressure deficit showed stomatal conductance dropping by only 25% in paper birch, while approaching zero in red oak and red maple. The study highlighted differences in ecophysiological controls on sap flux exerted by co-occurring species. Paper birch is a fast-growing, shade-intolerant species with an earlier successional status than red oak and red maple. Risking water loss through nighttime transpiration may provide paper birch with an ecological advantage by enabling the species to maximize photosynthesis and support rapid growth. Nighttime transpiration may also be a mechanism for delivering oxygen to respiring cells in the deep sapwood of paper birch.     

Spatial correlation of pit and mound topography with canopy gaps in a virgin mixed beech forest,northern Iran

In a reserved forest parcel in a virgin eastern Hyrcanian mixed beech forest,80 ha was surveyed to determine the pit and mound topography,canopy gaps and dead trees.The aim was to investigate the spatial patterns and correlation of pit and mound features with canopy gaps.Seventy-five canopy gaps and 61 pit and mound features were identified.The univariate first order nearest neighbor(R_CE)and bivariate second order test(Ripley’s K)statistic were applied.R_CE statistics highlighted a general aggregation pattern for canopy gaps and pits and mounds,while pits and mounds alone were more clumped.Distances between canopy gaps were 130 m average,whereas distances between pit and mound features and dead trees were 60 and 78 m,respectively.Spatial positive correlation of canopy gaps with pits and mounds were observed with all distances.The result of spatial correlations between canopy gaps with pits and mounds confirmed that windthrows cause micro successions in fallen tree ecosystem-scale correlated with gap phase dynamics in the forest community-scale.     

Dynamics of leaf area index and canopy openness of three forest types in a warm temperate zone

Deciduous broad-leaved forests (DBF), Larix principis-rupprechtii (LF) and Pinus tabulaeformis plantations (PF) are three typical forest communities in the warm temperate zone of the Dongling Mountains. In this study, we used an indirect method, hemispheric photography, to measure and analyze the dynamics of leaf area index (LAI) and canopy openness of the three forest communities. The results show that the LAI values of DBF and LF increased gradually with plant growth and development. The highest LAI value appeared in August, while canopy openness changed inversely with LAI. The lowest value appeared in November. DBF maintained a higher LAI in August and had a more open canopy in November compared with LF. For PF, we observed little changes in the LAI and canopy openness which was attributed to the leaf retention of this evergreen species. However, a similar relation between LAI and canopy openness was found for the three forest communities: canopy openness varied inversely with LAI. The relation is exponential and significant. Therefore, canopy openness is a good indicator of LAI in forests. This result can be used to test the validity of the LAI based on remote sensing and to provide a reference for the study of the canopy heterogeneity and its effect. This also benefits modeling for fluxes of carbon, water and energy from the level of the stand to landscape. __________ Translated from Journal of Plant Ecology, 2007, 31(3): 431–436 [译自: 植物生态学报]     

Interannual consistency in canopy stomatal conductance control of leaf water potential across seven tree species

We investigated interannual variability of canopy transpiration per unit ground area (E (C)) and per unit leaf area (E (L)) across seven tree species in northern Wisconsin over two years. These species have previously been shown to be sufficient to upscale stand-level transpiration to the landscape level during one growing season. Our objective was to test whether a simple plant hydraulic model could capture interannual variation in transpiration. Three species, wetland balsam fir (Abies balsamea (L.) Mill), basswood (Tilia Americana L.) and speckled alder (Alnus rugosa (DuRoi) Spreng), had no change in E (C) or E (L) between 2000 and 2001. Red pine (Pinus resinosa Ait) had a 57 and 19% increase in E (C) and E (L), respectively, and sugar maple (Acer saccharum Marsh) had an 83 and 41% increase in E (C) and E (L), respectively, from 2000 to 2001. Quaking aspen (Populus tremuloides Michx) had a 50 and 21% decrease in E (C) and E (L), respectively, from 2000 to 2001 in response to complete defoliation by forest tent caterpillar (Malascoma distria Hüber) and subsequent lower total leaf area index of the reflushed foliage. White cedar (Thuja occidentalis L.) had a 20% decrease in both E (C) and E (L) caused by lowered surface water in wetlands in 2001 because of lower precipitation and wetland flow management. Upland A. balsamea increased E (L) and E (C) by 55 and 53%, respectively, as a result of release from light competition of the defoliated, overstory P. tremuloides. We hypothesized that regardless of different drivers of interannual variability in E (C) and E (L), minimum leaf water potential would be regulated at the same value. Minimum midday water potentials were consistent over the two years within each of the seven species despite large changes in transpiration between years. This regulation was independently verified by the exponential saturation between daily E (C) and vapor pressure deficit (D) and the tradeoff between a reference canopy stomatal conductance (G (S)) and the sensitivity of G (S) to D, indicating that trees with high G (S) must decrease G (S) in response to atmospheric drought faster than trees with low G (S). Our results show that models of forest canopy transpiration can be simplified by incorporating G (S) regulation of minimum leaf water potential for isohydric species.     

Shoot growth responses to light microenvironment and correlative inhibition in tree seedlings under a forest canopy

To examine the mechanisms underlying crown development, I investigated the dependence of shoot behavior on light microenvironment in saplings of the evergreen broad-leaved tree species, Litsea acuminata (Bl.) Kurata, growing on a forest floor. The local light environment of individual shoots (shoot irradiance) and plants (plant irradiance, defined as the shoot irradiance of the most sunlit shoot of a plant) were analyzed as factors affecting shoot behavior. Daughter shoots that developed under partially sunlit conditions were longer and less leafy than daughter shoots developed under shaded conditions. Shoot production increased with increasing shoot irradiance. Terminal shoots receiving 5% or less of full sunlight produced 0.67 daughter shoots on average, whereas shoots receiving 10% or more of full sunlight produced 1.72 daughter shoots. In terminal shoots receiving 5% or less of full sunlight, the probability of producing no daughter shoots was about 63% when other shoots on the plant received 10% or more of full sunlight, but was < 35% where the rest of the plant was also shaded. Shoot death was observed only in shoots receiving 5% or less of full sunlight. The mortality of shaded shoots was higher in plants growing in high irradiance than in plants growing in low irradiance. The ecological significance of correlative inhibition (the enhanced mortality and reduced production of new shaded shoots in the presence of partially-sunlit shoots) is discussed.     

Comparative measurements of transpiration and canopy conductance in two mixed deciduous woodlands differing in structure and species composition

Transpiration of two heterogeneous broad-leaved woodlands in southern England was monitored by the sap flux technique throughout the 2006 growing season. Grimsbury Wood, which had a leaf area index (LAI) of 3.9, was dominated by oak (Quercus robur L.) and birch (Betula pubescens L.) and had a continuous hazel (Corylus avellana L.) understory. Wytham Woods, which had an LAI of 3.6, was dominated by ash (Fraxinus excelsior L.) and sycamore (Acer pseudoplatanus L.) and had only a sparse understory. Annual canopy transpiration was 367 mm for Grimsbury Wood and 397 mm for Wytham Woods. These values were similar to those for beech (Fagus sylvatica L.) plantations in the same region, and differ from one another by less than the typical margin of uncertainty of the sap flux technique. Canopy conductance (g(c)), calculated for both woodlands by inverting the Penman-Monteith equation, was related to incoming solar radiation (R(G)) and the vapor pressure deficit (D). The response of g(c) to R(G) was similar for both forests. Both reference conductance (g(cref)), defined as g(c) at D=1 kPa, and stomatal sensitivity (-m), defined as the slope of the logarithmic response curve of g(c) to D, increased during the growing season at Wytham Woods but not at Grimsbury Wood. The -m/g(cref) ratio was significantly lower at Wytham Woods than at Grimsbury Wood and was insufficient to keep the difference between leaf and soil water potentials constant, according to a simple hydraulic model. This meant that annual water consumption of the two woodlands was similar despite different regulatory mechanisms and associated short-term variations in canopy transpiration. The -m/g(cref) ratio depended on the range of D under which the measurements were made. This was shown to be particularly important for studies conducted under low and narrow ranges of D.     

Leaf hydraulic conductance in relation to anatomical and functional traits during Populus tremula leaf ontogeny

Leaf hydraulic conductance (K(leaf)) and several characteristics of hydraulic architecture and physiology were measured during the first 10 weeks of leaf ontogeny in Populus tremula L. saplings growing under control, mild water deficit or elevated temperature conditions. During the initial 3 weeks of leaf ontogeny, most measured characteristics rapidly increased. Thereafter, a gradual decrease in K(leaf) was correlated with a decrease in leaf osmotic potential under all conditions, and with increases in leaf dry mass per area and bulk modulus of elasticity under mild water deficit and control conditions. From about Week 3 onward, K(leaf) was 33% lower in trees subjected to mild water deficit and 33% higher in trees held at an elevated temperature relative to control trees. Mild water deficit and elevated temperature treatment had significant and opposite effects on most of the other characteristics measured. The ontogenetic maximum in K(leaf) was correlated positively with the width of xylem conduits in the midrib, but negatively with the overall width of the midrib xylem, number of lateral ribs, leaf dry mass per area and bulk modulus of elasticity. The ontogenetic maximum in K(leaf) was also correlated positively with the proportion of intercellular spaces and leaf osmotic potential, but negatively with leaf thickness, volume of mesophyll cells and epidermis and number of cells per total mesophyll cell volume, the closest relationships being between leaf osmotic potential and number of cells per total mesophyll cell volume. It was concluded that differences in protoplast traits are more important than differences in xylem or parenchymal cell wall traits in determining the variability in K(leaf) among leaves growing under different environmental conditions.     

Natural regeneration of oriental beech(Fagus orientalis Lipsky)trees in canopy gaps and under closed canopy in a forest in northern Iran

For developing nature-based silvicultural practices in a beech forest to promote, abundance, height,vitality, and preferred growth form, regenerated trees growing in gaps were compared with those under closed canopies.A systematic 50×50 m grid was plotted in a beech stand in the Kheyrud Experimental Forest for selecting trees to measure variables in gaps and under closed canopies.Abundance and mean height of regenerated beech trees were significantly higher in closed canopies than in canopy gaps.Beech seedlings with excurrent growth were significantly taller within regeneration patches under closed canopy.Moreover, regenerated trees with high vitality were more abundant in closed canopy areas than in gaps.Thus, beech regeneration should improve under closed canopies; hence, gaps in a near natural forest should be created only after adequate regeneration and appropriate growth under the parent tree in a closed canopy area is ensured.The results of this research have profound implications for the sustainable management of the forest and for ensuring sustainable beech regeneration.The presence of a closed canopy cover likely will reduce potential stresses on oriental beech regeneration.     

Effects of woodland structure on breeding bird populations in stands of coppiced lime (Tilia cordata) in western England over a 10-year period

Bird populations were estimated in three adjacent stands ofsmall-leaved lime (Tilia cordata) at Shrawley Wood, Worcestershire,between 1984 and 1993. The stands were old coppice, recent coppicecut in the late 1970s and thinned coppice that had been singledprobably in the 1970s. It is thought that the stands were allpreviously coppiced in the 1940s. The study area presented anunusual opportunity to (1) assess effects of woodland structureon bird populations in the absence of any major differencesin tree species, (2) to compare the dynamics of bird populations,and hence assess habitat preferences, within large samples ofeach habitat. Overall densities of birds were consistently lowestin the old coppice and generally highest in the recent coppice.The singled coppice supported the least diverse bird communitiesas measured by a simple diversity index and the expected numbersof species within standardized samples of individuals. Warblers(Sylviidae) were strikingly more abundant in recent coppicethan other habitats, being virtually absent from the old coppice.Wrens were by far the most abundant species in the singled coppice.There was strong evidence that the old coppice was the leastpreferred habitat of wren and robin. The most striking changeduring the study was the collapse of the warbler populationsin the recent coppice after some 11 to 12 years of coppice growth.This was predictable from previous studies, but the timing ofthe warbler decline occurred later than expected and was probablya consequence of the relatively slow canopy closure of the coppiceat Shrawley Wood. Differences between stands, and changes overthe 10 years within stands, were interpreted in terms of habitatstructure and effects of winter weather on survival of residentspecies. The findings are discussed in relation to managementof abandoned coppice; it is suggested that for conservationpurposes thinning should be regarded as a complementary approach,alongside coppicing and non-intervention.     

Light response of hydraulic conductance in bur oak (Quercus macrocarpa) leaves

A four- to seven-fold enhancement of leaf hydraulic conductance by light has been reported in three temperate tree species. The enhancement occurs in the liquid-flow pathway between the petiole and the site of water evaporation. The enhancement occurs within 1 h, and dissipates in darkness over a period of 1 to 10 h depending on species. Here we report light-induced enhancement of leaf hydraulic conductance in a fourth species, bur oak (Quercus macrocarpa Michx.), the dependence of the effect on light flux and color, its absence in leaves of seedlings, and the impact on the response of leaf vein severance and several metabolic inhibitors. The light response of leaf hydraulic conductance approached saturation at a photosynthetic photon flux of 150 mumol m(-2) s(-1). Hydraulic enhancement was greater in response to blue and green light than to visible radiation of longer wavelengths, although at the same irradiance, the response to white light was greater than to light of any single color. Atrazine (a photosystem II inhibitor), fusicoccin (which stimulates plasma membrane-bound H(+)-ATPase) and HgCl(2) (an aquaporin blocker) reduced the light response of leaf lamina hydraulic conductance. When 2-mercaptoethanol was added following mercury treatment, the light response was totally suppressed. Our results are consistent with the notion that the effect of light on leaf lamina hydraulic conductance is controlled by factors acting outside the leaf veins, possibly through light-induced changes in membrane permeability of either mesophyll or bundle sheath cells, or both.     

Effects of CO2 enrichment on the photosynthetic light response of sun and shade leaves of canopy sweetgum (Liquidambar styraciflua) in a forest ecosystem

To investigate whether sun and shade leaves respond differently to CO2 enrichment, we examined photosynthetic light response of sun and shade leaves in canopy sweetgum (Liquidambar styraciflua L.) trees growing at ambient and elevated (ambient + 200 microliters per liter) atmospheric CO2 in the Brookhaven National Laboratory/Duke University Free Air CO2 Enrichment (FACE) experiment. The sweetgum trees were naturally established in a 15-year-old forest dominated by loblolly pine (Pinus taeda L.). Measurements were made in early June and late August 1997 during the first full year of CO2 fumigation in the Duke Forest FACE experiment. Sun leaves had a 68% greater leaf mass per unit area, 63% more leaf N per unit leaf area, 27% more chlorophyll per unit leaf area and 77% greater light-saturated photosynthetic rates than shade leaves. Elevated CO2 strongly stimulated light-saturated photosynthetic rates of sun and shade leaves in June and August; however, the relative photosynthetic enhancement by elevated CO2 for sun leaves was more than double the relative enhancement of shade leaves. Elevated CO2 stimulated apparent quantum yield by 30%, but there was no interaction between CO2 and leaf position. Daytime leaf-level carbon gain extrapolated from photosynthetic light response curves indicated that sun leaves were enhanced 98% by elevated CO2, whereas shade leaves were enhanced 41%. Elevated CO2 did not significantly affect leaf N per unit area in sun or shade leaves during either measurement period. Thus, the greater CO2 enhancement of light-saturated photosynthesis in sun leaves than in shade leaves was probably a result of a greater amount of nitrogen per unit leaf area in sun leaves. A full understanding of the effects of increasing atmospheric CO2 concentrations on forest ecosystems must take account of the complex nature of the light environment through the canopy and how light interacts with CO2 to affect photosynthesis.