Leaf macro- and micro-morphological altitudinal variability of Carpinus betulus in the Hyrcanian forest (Iran)

We investigated the altitudinal variation of Carpinus betulus L. in the Hyrcanian forest using leaf macro-morphological and micro-morphological traits. We collected a total of 1600 leaves from two locations. In each location, we sampled six populations along an altitudinal gradient ranging from 100 m to 1,150 m. We found that trees in the higher elevations have smaller leaf lamina than those in the lower elevations. In contrast, leaf mass per area was high at low altitudes and increased newly at the higher ones. Stomatal dimension was negatively correlated with elevation, while stomatal density was positively correlated with elevation. We also found that two transects showed the same plasticity trend. Leaf area showed the highest plasticity, while the number of veins showed the lowest plasticity. This study shows that altitude, and related temperature and rainfall, represents an important driving force in Carpinus betulus leaf morphological variation. Moreover, our results suggest that leaf area, leaf mass per area and stomatal density could influence the species responses to different ecological conditions.     

Radiation-use efficiency of a forest exposed to elevated concentrations of atmospheric carbon dioxide

We compared radiation-use efficiency of growth (epsilon;), defined as rate of biomass accumulation per unit of absorbed photosynthetically active radiation, of forest plots exposed to ambient (approximately 360 micro l l-1) or elevated (approximately 560 micro l l-1) atmospheric CO2 concentration ([CO2]). Large plots (30-m diameter) in a loblolly pine (Pinus taeda L.) plantation, which contained several hardwood species in the understory, were fumigated with a free-air CO2 enrichment system. Biomass accumulation of the dominant loblolly pines was calculated from monthly measurements of tree growth and site-specific allometric equations. Depending on the species, leaf area index (L*) was estimated by three methods: optical, allometric and litterfall. Based on the relationship between tree height and diameter during the first 3 years of exposure, we conclude that elevated [CO2] did not alter the pattern of aboveground biomass allocation in loblolly pine. There was considerable variation in L* estimates by the different methods; total L* was 18-42% lower when estimated by the optical method compared with estimates from allometric calculations, and this discrepancy was reduced when optical measurements were corrected for the non-random distribution of loblolly pine foliage. The allometric + litterfall approach revealed a seasonal maximum total L* of 6.2-7.1 with about 1/3 of the total from hardwood foliage. Elevated [CO2] had only a slight effect on L* in the first 3 years of this study. Mean epsilon; (+/- SD), calculated for loblolly pine only, was 0.49 +/- 0.05 and 0.62 +/- 0.04 g MJ-1 for trees in the ambient and elevated [CO2] plots, respectively. The 27% increase in epsilon; in response to CO2 enrichment was caused primarily by the stimulation of biomass increment, as there was only a small effect of elevated [CO2] on L* during the initial years of fumigation. Long-term increases in atmospheric [CO2] can increase epsilon; in closed-canopy forests but the absolute magnitude and duration of this increase remain uncertain.     

Stand-level patterns of carbon fluxes and partitioning in a Eucalyptus grandis plantation across a gradient of productivity, in Sao Paulo State, Brazil

Wood production represents a large but variable fraction of gross primary production (GPP) in highly productive Eucalyptus plantations. Assessing patterns of carbon (C) partitioning (C flux as a fraction of GPP) between above- and belowground components is essential to understand mechanisms driving the C budget of these plantations. Better knowledge of fluxes and partitioning to woody and non-woody tissues in response to site characteristics and resource availability could provide opportunities to increase forest productivity. Our study aimed at investigating how C allocation varied within one apparently homogeneous 90 ha stand of Eucalyptus grandis (W. Hill ex Maiden) in Southeastern Brazil. We assessed annual above-ground net primary production (ANPP: stem, leaf, and branch production) and total belowground C flux (TBCF: the sum of root production and respiration and mycorrhizal production and respiration), GPP (computed as the sum of ANPP, TBCF and estimated aboveground respiration) on 12 plots representing the gradient of productivity found within the stand. The spatial heterogeneity of topography and associated soil attributes across the stand likely explained this fertility gradient. Component fluxes of GPP and C partitioning were found to vary among plots. Stem NPP ranged from 554 g C m(-2) year(-1) on the plot with lowest GPP to 923 g C m(-2) year(-1) on the plot with highest GPP. Total belowground carbon flux ranged from 497 to 1235 g C m(-2) year(-1) and showed no relationship with ANPP or GPP. Carbon partitioning to stem NPP increased from 0.19 to 0.23, showing a positive trend of increase with GPP (R(2) = 0.29, P = 0.07). Variations in stem wood production across the gradient of productivity observed at our experimental site were a result of the variability in C partitioning to different forest system components.     

Belowground competition in a broad-leaved temperate mixed forest: pattern analysis and experiments in a four-species stand

We investigated fine root biomass and distribution patterns in a species-rich temperate Carpinus–Quercus–Fagus–Tilia forest and searched for experimental evidence of symmetry or asymmetry in belowground competition. We conducted extensive root coring and applied the recently introduced in situ-root growth chamber technique for quantifying fine root growth under experimentally altered intra- and interspecific root neighbourhoods in the intact stand. In 75% of all soil cores, fine roots of more than two tree species were present indicating a broad overlap of the root systems of neighbouring trees. Quercus trees had more than ten times less fine root biomass in relation to aboveground biomass or productivity (stem growth) and a much higher leaf area index/root area index ratio than Carpinus, Fagus and Tilia trees. The root growth chamber experiments indicated a high belowground competitive ability of Fagus in interspecific interactions, but a low one of Quercus. We conclude that (1) interspecific root competition is ubiquitous in this mixed stand, (2) root competition between trees can be clearly asymmetric, and (3) tree species may be ranked according to their belowground competitive ability. Fagus was found to be the most successful species in belowground competition which matches with its superiority in aboveground competition in this forest community.     

Hydraulic conductance, light interception and needle nutrient concentration in Scots pine stands and their relations with net primary productivity

Aboveground xylem hydraulic conductance was determined in Scots pine (Pinus sylvestris L.) trees and stands from 7 to about 60 years of age. At the stand scale, leaf area index and net primary productivity (NPP, above- plus belowground) increased and reached a plateau at about 25-30 and 15-20 years, respectively; both parameters declined in mature stands. Stand hydraulic conductance followed a similar trend to NPP, with a maximum at about 15-20 years and a pronounced reduction in old stands. At the tree scale, annual biomass growth per unit of leaf area (growth efficiency) declined with tree age, whereas aboveground sapwood volume per unit leaf area, which is linearly related to maintenance respiration costs, steadily increased. Radiation interception per unit leaf area increased significantly with reduced leaf area index of mature stands, despite increased foliage clumping in the canopies of mature trees. Needle nutrient concentration did not change in the chronosequence. Tree hydraulic conductance per unit leaf area was strongly and positively correlated with growth efficiency. We discuss our findings in the context of growth reductions in mature and old trees, and suggest that increased hydraulic resistance and maintenance respiration costs may be the main causes of reduced carbon gain in mature and old trees.     

Do changes in carbon allocation account for the growth response to potassium and sodium applications in tropical Eucalyptus plantations?

Understanding the underlying mechanisms that account for the impact of potassium (K) fertilization and its replacement by sodium (Na) on tree growth is key to improving the management of forest plantations that are expanding over weathered tropical soils with low amounts of exchangeable bases. A complete randomized block design was planted with Eucalyptus grandis (W. Hill ex Maiden) to quantify growth, carbon uptake and carbon partitioning using a carbon budget approach. A combination of approaches including the establishment of allometric relationships over the whole rotation and measurements of soil CO(2) efflux and aboveground litterfall at the end of the rotation were used to estimate aboveground net production (ANPP), total belowground carbon flux and gross primary production (GPP). The stable carbon isotope (δ(13)C) of stem wood α-cellulose produced every year was used as a proxy for stomatal limitation of photosynthesis. Potassium fertilization increased GPP and decreased the fraction of carbon allocated belowground. Aboveground net production was strongly enhanced, and because leaf lifespan increased, leaf biomass was enhanced without any change in leaf production, and wood production (P(W)) was dramatically increased. Sodium application decreased the fraction of carbon allocated belowground in a similar way, and enhanced GPP, ANPP and P(W), but to a lesser extent compared with K fertilization. Neither K nor Na affected δ(13)C of stem wood α-cellulose, suggesting that water-use efficiency was the same among the treatments and that the inferred increase in leaf photosynthesis was not only related to a higher stomatal conductance. We concluded that the response to K fertilization and Na addition on P(W) resulted from drastic changes in carbon allocation.     

Physiological and phenological responses of oak seedlings to oak forest soil in the absence of trees

Established trees influence the growth and physiology of seedlings by altering above- and belowground conditions; however, tree influences on seedling physiology via belowground interactions are not well understood. We used soil transfers to an open field to examine the belowground influences of a Quercus ellipsoidalis E.J.Hill dominated forest on Q. ellipsoidalis seedling mycorrhizal infection, nutrient uptake, growth and photosynthesis over three years. After two years, seedlings planted with large quantities of forest soil (HF treatment) had greater leaf mass and foliar N concentrations than seedlings receiving smaller quantities of forest soil (LF) and control treatments. Mycorrhizal infection was greater in the HF treatment after one year compared with the LF and control treatments, with a positive correlation of foliar N and mycorrhizal infection in Year 2. There were marked effects of treatments on seedling spring phenology with HF seedlings breaking bud up to 17 days earlier than seedlings in the other treatments. The HF seedlings also had more rapid leaf expansion and larger leaves, and an increase in net photosynthetic rates. These results highlight complex linkages between above- and belowground physiology: forest soil had substantial effects on seedling physiology, including traits such as phenology that have previously been considered to be under aboveground control. Belowground influences of trees on conspecific seedlings may play a critical role in early seedling establishment.     

Ecophysiology of Cecropia schreberiana saplings in two wind regimes in an elfin cloud forest: growth,gas exchange,architecture and stem biomechanics

The hypothesis that the short, stunted development of elfin cloud forest trees is influenced by strong winds was tested by studying growth, gas exchange and biomechanics of potted saplings of Cecropia schreberiana Miq. ex. C. peltata L. (Cecropiaceae) exposed to two natural wind regimes (exposed and protected) at high elevation sites in the Luquillo Experimental Forest, Puerto Rico. The wind-exposure treatment produced several thigmomorphogenetic responses, including reductions in plant stature and crown area, changes in allocation patterns, and increased root to shoot ratio, leaf abrasion and leaf epinasty. Wind-exposure decreased maximum photosynthetic rate and respiration on an area basis, but not on a leaf-mass basis. Wind-exposed plants had lower apparent quantum yields, and higher light compensation points than wind-protected plants. Photosynthetic nitrogen-use efficiency was lower in wind-exposed plants, but such plants had higher leaf nitrogen concentration than wind-protected plants. There were no effects of treatments on stomatal conductance, transpiration rate and water-use efficiency. Stems of wind-exposed plants had lower second moment of area, apparent modulus of elasticity, flexural stiffness and stem density, but higher water content than wind-protected plants. Tissue-density-specific stiffness and the calculated critical height were not affected by the treatments. Wind-exposed plants were biomechanically less predisposed to bending and failing under their own weight than wind-protected plants because their safety factors were smaller, indicating that maintenance of an ontogenetically less developed structure enables plants to cope with wind loading. Windward trees showed a lower scaling component of the allometric relationship between diameter and height than leeward trees.     

Quantitative relationships between fine roots and stand characteristics

Fine roots absorb nutrients and water for photosynthesizing leaves, which in return provide them with hydrocarbon products. Knowledge of the fine root biomass (FRB) at the individual tree level and its relationships with other components related to tree growth, especially leaves aboveground, is scarce. Therefore, we reviewed the FRB of major forest-forming species using a database of 518 forest stands compiled from the literature, including 21 tree species and 16 shrub species, in order to confirm the relationships between environmental or forest stand variables and FRB at the stand and tree levels, and we further determine the relationships between fine roots belowground and leaves aboveground. Correlations between FRB and site characteristics (latitude, elevation, age, density, and basal area) appeared to be species-specific. There were hardly any significant correlations between stand FRB and latitude, elevation, stand age and stand density. Tree FRB was better correlated with tree basal area than stand FRB with stand basal area. There was a significant linear relationship between tree FRB and tree basal area. In addition, individual FRB was significantly linearly related to leaf biomass for all analyzed species. When these species were grouped into coniferous and deciduous, or all species together, there were still significant linear relationships between tree FRB and tree basal area and leaf biomass. The ratios of FRB to leaf biomass varied between and among species and even among regions for the same species. For both Picea abies and Pinus sylvestris, the ratio of FRB to leaf biomass was negatively related to the ratio of annual actual evapotranspiration to annual potential evapotranspiration, which was an indicator of water availability.     

Radiation use efficiency in adjacent hardwood and pine forests in the southern Appalachians

The efficiency with which trees convert photosynthetically active radiation (PAR) to biomass has been shown to be consistent within stands of an individual species, which is useful for estimating biomass production and carbon accumulation. However, radiation use efficiency (?) has rarely been measured in mixed-species forests, and it is unclear how species diversity may affect the consistency of ?, particularly across environmental gradients. We compared aboveground net primary productivity (ANPP), intercepted photosynthetically active solar radiation (IPAR), and radiation use efficiency (? = ANPP/IPAR) between a mixed deciduous forest and a 50-year-old white pine (Pinus strobus L.) plantation in the southern Appalachian Mountains. Average ANPP was similar in the deciduous forest (11.5 Mg ha⁻¹ y⁻¹) and pine plantation (10.2 Mg ha⁻¹ y⁻¹), while ? was significantly greater in the deciduous forest (1.25 g MJ⁻¹) than in the white pine plantation (0.63 g MJ⁻¹). Our results demonstrate that late-secondary hardwood forests can attain similar ANPP as mature P. strobus plantations in the southern Appalachians, despite substantially less annual IPAR and mineral-nitrogen availability, suggesting greater resource-use efficiency and potential for long-term carbon accumulation in biomass. Along a 260 m elevation gradient within each forest there was not significant variation in ?. Radiation use efficiency may be stable for specific forest types across a range of environmental conditions in the southern Appalachian Mountains, and thus useful for generating estimates of ANPP at the scale of individual watersheds.     

Production physiology of three fast-growing hardwood species along a soil resource gradient

We determined how specific leaf area (SLA), specific leaf nitrogen (SLN), leaf area index (LAI), light-saturated photosynthesis (Amax) and aboveground net primary productivity (ANPP) of three commercially important hardwood species, eastern cottonwood (Populus deltoides Bartr.), American sycamore (Platanus occidentalis L.) and cherrybark oak (Quercus falcata var.pagodafolia Ell.), vary across a soil resource gradient. Five treatments were applied in a randomized block design (control, irrigation only (IRR), and irrigation plus fertilization with 56, 112 or 224 kg N ha-1 year-1 (N56, N112 and N224)) with four replications per species. When trees were 6 years old, Amax, SLA, SLN, LAI and ANPP were quantified during peak leaf production within a single growing season. In all species, Amax for sun leaves was significantly higher than for shade leaves (34, 32 and 29 micromol m2 s-1 versus 27, 23 and 23 micromol m2 s-1 for cottonwood, cherrybark oak and sycamore sun and shade leaves, respectively) and tended to plateau in the N112 treatment. The SLA was significantly lower in sun than in shade leaves and reached a plateau in IRR-treated cottonwood and sycamore, and in N56-treated oak. Values of SLN peaked in the N122 treatment for cottonwood sun leaves (1.73 g N m2) and in the N56 treatment for sycamore and oak (1.54 and 1.90 g N m2, respectively). In sun and shade leaves of all species, Amax increased with increasing SLN. Cherrybark oak LAI reached a plateau across the resource gradient in the N56 treatment, whereas cottonwood and sycamore LAI reached a plateau in the IRR treatment. All species exhibited significant curvilinear relationships between canopy Amax and ANPP. These findings indicate that nutrients and water regulate leaf-level traits such as SLA and SLN, which in turn influence LAI and canopy photosynthesis, thereby affecting ANPP at the tree and stand levels.     

Drought sensitivity of Norway spruce is higher than that of silver fir along an altitudinal gradient in southwestern Germany

Context

For Central Europe, climate projections foresee an increase in temperature combined with decreasing summer precipitation, resulting in drier conditions during the growing season. This might negatively affect forest growth, especially at sites that are already water-limited, i.e., at low elevation. At higher altitudes trees might profit from increasing temperatures.

Aims

We analyzed variations in radial growth of silver fir (Abies alba Mill.) and Norway spruce (Picea abies (L.) Karst.) along an altitudinal gradient from 400 until 1,140 m a.s.l. in the Black Forest, to assess climate responses with increasing elevation.

Methods

Climate–growth relationships were analyzed retrospectively using tree-ring and climate data. In total, we sampled stem discs of 135 trees to build 27 species- and site-specific chronologies (n _fir?=?13, n _spruce?=?14).

Results

Our results indicate distinct differences in climate–growth relations between fir and spruce along the gradient. Growth of high-altitude fir was positively related to temperature from January till March. Growth of low-altitude fir and spruce at all elevations was positively related to precipitation and negatively to temperature during the growing season, particularly in July. A self-calibrating Palmer drought severity index underlined summer drought sensitivity of these trees.

Conclusion

Overall, we found that climatic control of tree growth changes over altitude for fir. For spruce, a remarkable synchrony in growth variation and climate response was shown, which indicates that this species is drought sensitive at all studied elevations. In a future warmer climate, the growth of low-altitude fir and spruce along the entire studied gradient may be negatively affected in the Black Forest, if an increased evaporative demand cannot be compensated by increased water supply.     

Does initial spacing influence crown and hydraulic architecture of Eucalyptus marginata?

Long-term declines in rainfall in south-western Australia have resulted in increased interest in the hydraulic characteristics of jarrah (Eucalyptus marginata Donn ex Smith) forest established in the region's drinking water catchments on rehabilitated bauxite mining sites. We hypothesized that in jarrah forest established on rehabilitated mine sites: (1) leaf area index (L) is independent of initial tree spacing; and (2) more densely planted trees have less leaf area for the same leaf mass, or the same sapwood area, and have denser sapwood. Initial stand densities ranged from about 600 to 9000 stems ha(-1), and trees were 18 years old at the time of sampling. Leaf area index was unaffected by initial stand density, except in the most sparsely stocked stands where L was 1.2 compared with 2.0-2.5 in stands at other spacings. The ratio of leaf area to sapwood area (A(l):A(s)) was unaffected by tree spacing or tree size and was 0.2 at 1.3 m height and 0.25 at the crown base. There were small increases in sapwood density and decreases in leaf specific area with increased spacing. Tree diameter or basal area was a better predictor of leaf area than sapwood area. At the stand scale, basal area was a good predictor of L (r(2) = 0.98, n = 15) except in the densest stands. We conclude that the hydraulic attributes of this forest type are largely independent of initial tree spacing, thus simplifying parameterization of stand and catchment water balance models.     

Edge effects and forest water use: A field study in a mixed deciduous woodland

A field study was carried out in a mixed deciduous forest in order to measure the spatial variability of evapotranspiration in relation to distance from the nearest forest edge. Throughfall was collected in storage gauges in a transect across the edge. Transpiration was measured at the tree scale by means of the sap flux technique. Thermal dissipation probes were inserted into the hydro-active sapwood of 12–16 sample trees at a time covering four species. The sample trees were located close to a north- and a south-facing forest edge and between 3 and 69 m away from the nearest edge. The probes were moved to new trees about once a month and in total 71 trees were sampled. Sap flux densities were compared with potential evaporation and scaled up to the stand through multiplication with sapwood area per unit ground area. No significant edge effect on interception evaporation could be detected but there was a large influence on stand transpiration which increased towards the edge. In ash (Fraxinus excelsior L.), this increase resulted mainly from enhanced sap flux density (by 33–82%, depending on the size class) in trees located at the edge, whereas in oak (Quercus robur L.) the sap flux density was similar in edge and inner trees and an effect was only found at the stand scale in the way that the total basal area, per unit ground area, was larger near the forest edge than in the forest interior. Hawthorn (Crataegus monogyna L.) and field maple (Acer campestre L.), which occurred mainly in the understorey, were only weakly affected by the proximity to an edge. At the stand scale the total seasonal transpiration varied between 354 mm in the forest interior (>45 m away from the edge) and 565 mm at the forest edge (<15 m away from the edge), whilst the potential evaporation over the same period was 571 mm. This corresponds to Priestley–Taylor coefficients of 0.78 in the interior and 1.25 at the edge, whilst intermediate numbers were found for the area between the edge and inner zones. Using these results to calculate the average water loss per unit ground area of hypothetical woodlands of various sizes, it is shown that the edge effect dominates the water use of small forests and becomes negligible only for woodlands larger than 100 ha.     

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.     

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.     

Plant size, not age, regulates growth and gas exchange in grafted Scots pine trees

We studied the effect of scion donor-tree age on the physiology and growth of 6- to 7-year-old grafted Scots pine (Pinus sylvestris L.) trees (4 and 5 years after grafting). Physiological measurements included photosynthethetic rate, stomatal conductance, transpiration, whole plant hydraulic conductance, needle nitrogen concentration and carbon isotope composition. Growth measurements included total and component biomasses, relative growth rates and growth efficiency. Scion donor trees ranged in age from 36 to 269 years at the time of grafting. Hydraulic conductance was measured gravimetrically, applying the Ohm's law analogy, and directly, with a high-pressure flow meter. We found no effect of scion donor-tree age on any of the variables measured. There was, however, great variation within scion donor-tree age groups, which was related to the size of the grafted trees. Differences in size may have been caused by variable initial grafting success, but there was no indication that grafting success and age were related. At the stem level, hydraulic conductance scaled with total leaf area so that total conductance per unit leaf area did not vary with crown size. However, leaf specific hydraulic conductance (gravimetric), transpiration, photosynthesis and stomatal conductance declined with increasing total tree leaf area and needle width. We hypothesize that needle width is inversely related to mesophyll conductance. We conclude that canopy and needle size and not scion donor-tree age determined gas exchange in our grafted trees.     

Transpiration and forest structure in relation to soil waterlogging in a Hawaiian montane cloud forest

Transpiration, leaf characteristics and forest structure in Metrosideros polymorpha Gaud. stands growing in East Maui, Hawaii were investigated to assess physiological limitations associated with flooding as a mechanism of reduced canopy leaf area in waterlogged sites. Whole-tree sap flow, stomatal conductance, microclimate, soil oxidation-reduction potential, stand basal area and leaf area index (LAI) were measured on moderately sloped, drained sites with closed canopies (90%) and on level, waterlogged sites with open canopies (50-60%). The LAI was measured with a new technique based on enlarged photographs of individual tree crowns and allometric relationships. Sap flow was scaled to the stand level by multiplying basal area-normalized sap flow by stand basal area. Level sites had lower soil redox potentials, lower mean stand basal area, lower LAI, and a higher degree of soil avoidance by roots than sloped sites. Foliar nutrients and leaf mass per area (LMA) in M. polymorpha were similar between level and sloped sites. Stomatal conductance was similar for M. polymorpha saplings on both sites, but decreased with increasing tree height (r(2) = 0.72; P < 0.001). Stand transpiration estimates ranged from 79 to 89% of potential evapotranspiration (PET) for sloped sites and from 28 to 51% of PET for level sites. Stand transpiration estimates were strongly correlated with LAI (r(2) = 0.96; P < 0.001). Whole-tree transpiration was lower at level sites with waterlogged soils, but was similar or higher for trees on level sites when normalized by leaf area. Trees on level sites had a smaller leaf area per stem diameter than trees on sloped sites, suggesting that soil oxygen deficiency may reduce leaf area. However, transpiration per unit leaf area did not vary substantially, so leaf-level physiological behavior was conserved, regardless of differences in tree leaf area.     

Effects of varying crop load on photosynthesis, dry matter production and partitioning of Crispin/M.27 apple trees

Fruit load was altered by flower thinning on three- and four-year-old, field-grown apple trees. Increasing fruit load led to increases in dry matter production per unit leaf area and partitioning to fruit and to decreases in fruit size, percentage fruit dry matter, dry matter partitioning to new shoot growth, thickening of existing woody tissue and root growth. Flower bud production for the following spring was also negatively affected by an increase in fruit load. Leaf photosynthesis was increased in cropping trees in July and August at the time of maximum fruit dry weight increase. Calculated light interception was linearly related to leaf area. The efficiency of conversion of intercepted photosynthetic active radiation to dry matter energy equivalents was 3.3% in heavily cropping trees and 1.8% in non-cropping trees. Total dry matter production was linearly related to both leaf area and light interception, but the variance accounted for by the regression was more than doubled if fruit dry matter or fruit number was included in the regression.