An investigation of hydraulic limitation and compensation in large,old Douglas-fir trees

The hydraulic limitation hypothesis (Ryan and Yoder 1997) proposes that leaf-specific hydraulic conductance (kl) and stomatal conductance (gs) decline as trees grow taller, resulting in decreased carbon assimilation. We tested the hydraulic limitation hypothesis by comparison of canopy-dominant Douglas-fir (Pseudotsuga menziesii var. menziesii (Mirb.) Franco) trees in stands that were approximately 15 m (20 years old), 32 m (40 years old) and 60 m (> 450 years old) tall in Wind River, Washington, USA. Carbon isotope discrimination (Delta) declined with tree height (18.6, 17.6 and 15.9 per thousand for stands 15, 32 and 60 m tall, respectively) indicating that gs may have declined proportionally with tree height in the spring months, when carbon used in the construction of new foliage is assimilated. Hydraulic conductance decreased by 44% as tree height increased from 15 to > 32 m, and showed a further decline of 6% with increasing height. The general nonlinear pattern of kl versus height was predicted by a model based on Darcy's Law. Stemwood growth efficiency also declined nonlinearly with height (60, 35 and 28 g C m-2 leaf area for the 15-, 32- and 60-m stands, respectively). Unlike kl and growth efficiency, gs and photosynthesis (A) during summer drought did not decrease with height. The lack of decline in cuvette-based A indicates that reduced A, at least during summer months, is not responsible for the decline in growth efficiency. The difference between the trend in gs and A and that in kl and D may indicate temporal changes (spring versus summer) in the response of gas exchange to height-related changes in kl or it may be a result of measurement inadequacies. The formal hydraulic limitation hypothesis was not supported by our mid-summer gs and A data. Future tests of the hydraulic limitation hypothesis in this forest should be conducted in the spring months, when carbon uptake is greatest. We used a model based on Darcy's Law to quantify the extent to which compensating mechanisms buffer hydraulic limitations to gas exchange. Sensitivity analyses indicated that without the observed increases in the soil-to-leaf water potential differential (DeltaPsi) and decreases in the leaf area/sapwood area ratio, kl would have been reduced by more than 70% in the 60-m trees compared with the 15-m trees, instead of the observed decrease of 44%. However, compensation may have a cost; for example, the greater DeltaPsi of the largest trees was associated with smaller tracheid diameters and increased sapwood cavitation, which may have a negative feedback on kl and gs.     

Evidence that hydraulic conductance limits photosynthesis in old Pinus ponderosa trees

We tested the hypotheses that hydraulic conductance is lower in old (about 250 years old and 30 m tall) compared to young (about 40 years old and 10 m tall) Pinus ponderosa Dougl. ex Laws. trees and that lower hydraulic conductance of old trees limits their photosynthesis. Hydraulic conductance at the end of summer 1995, calculated from leaf water potential and leaf gas exchange measurements on one-year-old needles, was 44% lower in old trees compared to young trees growing in a mixed age-class stand on the east slope of the Oregon Cascades. Whole-tree sapflow per unit leaf area averaged 53% lower in old trees compared to young trees and mean hydraulic conductance calculated from sapflow and water potential data was 63% lower in old trees than in young trees. For the entire summer, stomatal conductance (g(s)) and assimilation (A) declined more steeply with air saturation deficit (D) in old trees than in young trees. For both old and young trees, mean g(s) and A were approximately 32 and 21% lower, respectively, at typical midday D values (2.5-3.0 kPa). We hypothesized that if hydraulic conductance limits g(s) and A, then increasing or decreasing the leaf specific conductance of a branch will result in proportional changes in the responses of g(s) and A with D. Removal of 50% of the foliage from a set of experimental branches on old trees caused g(s) and A to decline less steeply with D in early summer, but values were not significantly different from control values in late summer. Cutting transverse notches in branches on young trees had no effect on the responses of g(s) and A with D. Leaf nitrogen content and photosynthetic capacity were similar suggesting that differences in g(s) and A between old and young trees were not caused by differences in photosynthetic capacity.     

Stomatal control and hydraulic conductance, with special reference to tall trees

A better understanding of the mechanistic basis of stomatal control is necessary to understand why modes of stomatal response differ among individual trees, and to improve the theoretical foundation for predictive models and manipulative experiments. Current understanding of the mechanistic basis of stomatal control is reviewed here and discussed in relation to the plant hydraulic system. Analysis focused on: (1) the relative role of hydraulic conductance in the vicinity of the stomatal apparatus versus whole-plant hydraulic conductance; (2) the influence of guard cell inflation characteristics and the mechanical interaction between guard cells and epidermal cells; and (3) the system requirements for moderate versus dramatic reductions in stomatal conductance with increasing evaporation potential. Special consideration was given to the potential effect of changes in hydraulic properties as trees grow taller. Stomatal control of leaf gas exchange is coupled to the entire plant hydraulic system and the basis of this coupling is the interdependence of guard cell water potential and transpiration rate. This hydraulic feedback loop is always present, but its dynamic properties may be altered by growth or cavitation-induced changes in hydraulic conductance, and may vary with genetically related differences in hydraulic conductances. Mechanistic models should include this feedback loop. Plants vary in their ability to control transpiration rate sufficiently to maintain constant leaf water potential. Limited control may be achieved through the hydraulic feedback loop alone, but for tighter control, an additional element linking transpiration rate to guard cell osmotic pressure may be needed.     

Fertilization effects on mean stomatal conductance are mediated through changes in the hydraulic attributes of mature Norway spruce trees

Stomatal conductance was quantified with sap flux sensors and whole-tree chambers in mature Norway spruce (Picea abies (L.) Karst.) trees after 3 years of exposure to elevated CO(2) concentration ([CO(2)]) in a 13-year nutrient optimization experiment. The long-term nutrient optimization treatment increased tree height by 3.7 m (67%) and basal diameter by 8 cm (68%); the short-term elevated [CO(2)] exposure had no effect on tree size or allometry. Nighttime transpiration was estimated as approximately 7% of daily transpiration in unchambered trees; accounting for the effect of nighttime flux on the processing of sap flux signals increased estimated daily water uptake by approximately 30%. Crown averaged stomatal conductance (g(s)) was described by a Jarvis-type model. The addition of a stomatal response time constant (tau) and total capacitance of stored water (C(tot)) improved the fit of the model. Model estimates for C(tot) scaled with sapwood volume of the bole in fertilized trees. Hydraulic support-defined as a lumped variable of leaf-specific hydraulic conductivity and water potential gradient (K(l)DeltaPsi) -was estimated from height, sapwood-to-leaf area ratio (A(s):A(l)) and changes in tracheid dimensions. Hydraulic support explained 55% of the variation in g(s) at reference conditions for trees across nutrient and [CO(2)] treatments. Removal of approximately 50% of A(l) from three trees yielded results suggesting that stomatal compensation (i.e., an increase in g(s)) after pruning scales inversely with K(l)DeltaPsi, indicating that the higher the potential hydraulic support after pruning, the less complete the stomatal compensation for the increase in A(s):A(l).     

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.     

Use of water stress integral to evaluate relationships between soil moisture,plant water stress and stand productivity in young Douglas-fir trees

Water stress integral (WSI) is a method of assessing cumulative plant water stress over a chosen period of time. While the technique has been used in other tree species, it has not been applied for reforestation projects. In this study we used the WSI approach for newly planted Douglas-fir in the Pacific Northwest (USA), where the Mediterranean climate, plant community development, and competition for water all play key roles in the success of establishment efforts. In this study, previously reported seedling growth, xylem water potential, and soil moisture data were utilized to provide direct correlations between Douglas-fir productivity, soil water availability and WSI. For each growing season, a strong relationship between WSI and volume growth as well as a strong linear relationship between WSI and soil moisture measured during mid-August was found. On average, for each reduction of 0.01 cm³ cm^?3 in soil moisture measured during mid-August, Douglas-fir seedling volume growth decreased by 5.6 and 7.7% in the first and second growing seasons, respectively. Preserving soil moisture until early-August through the judicial application of vegetation management regimes was critical for maximizing stand productivity. Based on these results, a single evaluation of soil volumetric water content during early-August can be used as a predictor of stand productivity during the initial two seasons of forest establishment.     

Canopy stomatal conductance and xylem sap abscisic acid (ABA) in mature Scots pine during a gradually imposed drought

We investigated the effect of drought on canopy stomatal conductance (g(c)), and examined the hypothesis that g(c) is controlled by the chemical messenger abscisic acid (ABA) produced in roots. Beginning in November 1994, we subjected a mature stand of Scots pine (Pinus sylvestris L.) to an imposed 11-month drought. Control plots were maintained at average-season soil water content. Xylem sap was extracted from shoots at regular intervals from April to November 1995. Soil water, sap flow and leaf water potentials (predawn to dusk) were recorded at the same time. Canopy stomatal conductance was calculated from sap flow data and xylem sap ABA concentrations ([ABA(xyl)]) were measured by radioimmunoassay. Mean [ABA(xyl)] in control trees was 250 micromol m(-3). No diurnal variation in [ABA(xyl)] was detected. With soil drying, [ABA(xyl)] increased to a maximum in summer (600 micromol m(-3)), but decreased again toward autumn; however, no significant increase in ABA flux to the leaves occurred. A decline in g(c) was detected when volumetric soil water content declined below 0.12. The decline in g(c) could not have been mediated by increasing [ABA(xyl)] because stomatal closure appeared to precede any increase in [ABA(xyl)]. Peak sap flow velocity data were used to estimate delivery times for root-to-shoot signals in 15-m tall trees. Under normal field conditions, a signal would take 12 days to travel from the site of production (roots) to the presumed site of action (shoots). However, under drought conditions it may take a chemical signal in excess of 6 weeks. We conclude that a feedforward model of short-term stomatal response to soil drying, based solely on the action of a chemical messenger from the roots, is not applicable in mature conifer trees because signal transmission is too slow.     

A potential role for xylem-phloem interactions in the hydraulic architecture of trees: effects of phloem girdling on xylem hydraulic conductance

We investigated phloem-xylem interactions in Acer rubrum L. and Acer saccharum Marsh. Our experimental method allowed us to determine xylem conductance of an intact branch by measuring the flow rate of water supplied at two delivery pressures to the cut end of a small side branch. We found that removal of bark tissue (phloem girdling) upstream of the point at which deionized water was delivered to the branch resulted in a decrease (24% for A. rubrum and 15% for A. saccharum) in branch xylem hydraulic conductance. Declines in hydraulic conductance with girdling were accompanied by a decrease in the osmotic concentration of xylem sap. The decrease in xylem sap concentration following phloem girdling suggests that ion redistribution from the phloem was responsible for the observed decline in hydraulic conductance. When the same measurements were made on branches perfused with KCl solution (approximately 140 mOsm kg(-1)), phloem girdling had no effect on xylem hydraulic conductance. These results suggest a functional link between phloem and xylem hydraulic systems that is mediated by changes in the ionic content of the cell sap.     

Transpiration-induced axial and radial tension gradients in trunks of Douglas-fir trees

We determined the axial and radial xylem tension gradients in trunks of young Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) trees. Axial specific conductivity (k(s-a)) and sap flux density (Js) were measured at four consecutive depths within the sapwood at a stem height of 1 m. By definition, at a given position in the bole, Js is a function not only of k(s-a) but also of the driving force for water movement. The Js:k(s-a) ratio was therefore used to estimate axial tension gradients and the radial gradients at a stem height of 1 m were calculated from the differences in axial tension gradients at each depth. Tracheid lumen diameter and tracheid length were used to predict differences in k(s-a) and its divergence from the theoretical k(s-a) determined by the Hagen Poisseuille equation. The ratio of k(s-a) (determined in the laboratory) to Js (measured in the field) varied with depth in the sapwood, resulting in non-uniform axial and radial tension gradients from inner to outer sapwood. Transpiration-induced axial tension gradients were in the range of 0.006-0.01 MPa m(-1) excluding the gravitational tension gradient. At a stem height of 1 m, radial tension gradients were in the range of 0.15-0.25 MPa m(-1) and were lower in the middle sapwood than in the inner or outer sapwood. Axial tension gradients were 44-50% higher in the outer sapwood than in the inner sapwood. At a stem height of 1 m, radial Js, calculated on the basis of radial tension gradients and measured radial specific conductivity (k(s-r)), was about two orders of magnitude smaller than axial Js. Our findings indicate that large radial tension gradients occur in the sapwood and clarify the role played by xylem k(s-a) and k(s-r) in determining in situ partitioning of Js in the axial and radial directions.     

Rate of stomatal opening, shoot hydraulic conductance and photosynthetic characteristics in relation to leaf abscisic acid concentration in six temperate deciduous trees

Correlations between leaf abscisic acid concentration ([ABA]), stomatal conductance (gs), rate of stomatal opening in response to an increase in leaf water potential (si), shoot hydraulic conductance (L) and photosynthetic characteristics were examined in saplings of six temperate deciduous tree species: Acer platanoides L., Padus avium Mill., Populus tremula L., Quercus robur L., Salix caprea L. and Tilia cordata Mill. Species-specific values of foliar [ABA] were negatively related to the mean values of gs, si, L and light- and CO2- saturated net photosynthesis (P(max)), thus providing strong correlative evidence of a scaling of foliar gas exchange and hydraulic characteristics with leaf endogenous [ABA]. In addition, we suggest that mean gs, si, L and Pmax for mature leaves may partly be determined by the species-specific [ABA] during leaf growth. The most drought-intolerant species had the lowest [ABA] and the highest gs, suggesting that interspecific differences in [ABA] may be linked to differences in species-specific water-use efficiency. Application of high concentrations of exogenous ABA led to large decreases in gs, si and P(max), further underscoring the direct role of ABA in regulating stomatal opening and photosynthetic rate. Exogenous ABA also decreased L, but the decreases were considerably smaller than the decreases in gs, si and Pmax. Thus, exogenous ABA predominantly affected the stomata directly, but modification of L by ABA may also be an important mechanism of ABA action. We conclude that interspecific variability in endogenous [ABA] during foliage growth and in mature leaves provides an important factor explaining observed differences in L, gs, si and Pmax among temperate deciduous tree species.     

Variation in water potential, hydraulic characteristics and water source use in montane Douglas-fir and lodgepole pine trees in southwestern Alberta and consequences for seasonal changes in photosynthetic capacity

Tree species response to climate change-induced shifts in the hydrological cycle depends on many physiological traits, particularly variation in water relations characteristics. We evaluated differences in shoot water potential, vulnerability of branches to reductions in hydraulic conductivity, and water source use between Pinus contorta Dougl. ex Loud. var. latifolia Engelm. (lodgepole pine) and Pseudotsuga menziesii (Mirb.) Franco (interior Douglas-fir), and determined the consequences for seasonal changes in photosynthetic capacity. The Douglas-fir site had soil with greater depth, finer texture and higher organic matter content than soil at the lodgepole pine site, all factors that increased the storage of soil moisture. While the measured xylem vulnerability curves were quite similar for the two species, Douglas-fir had lower average midday shoot water potentials than did lodgepole pine. This implied that lodgepole pine exhibited stronger stomatal control of transpiration than Douglas-fir, which helped to reduce the magnitude of the water potential gradient required to access water from drying soil. Stable hydrogen isotope measurements indicated that Douglas-fir increased the use of groundwater during mid-summer when precipitation inputs were low, while lodgepole pine did not. There was a greater reduction of photosynthetic carbon gain in lodgepole pine compared with Douglas-fir when the two tree species were exposed to seasonal declines in soil water content. The contrasting patterns of seasonal variation in photosynthetic capacity observed for the two species were a combined result of differences in soil characteristics at the separate sites and the inherent physiological differences between the species.     

Individual-tree height-, diameter- and crown-width increment equations for young Douglas-fir plantations

This work presents the results from the initial model development of a simulator to predict vegetation dynamics in young plantations growing in a Mediterranean environment. The simulator can predict growth dynamics for coniferous crop trees as well as competing hardwoods and shrubs. Model specification included conifer, shrub, and hardwood competition expressed at the plot-level. The system employs water-holding capacity as an indicator of productivity. Growth data were obtained from 109 plantations, ranging in age from 3 to 25, in southern Oregon and northern California. Douglas-fir (Pseudotsuga menziesii Mirb. Franco), the most common conifer species, was observed in 80 of the sampled stands. These observations were used for model development of the primary driving functions, which forecast 2-year basal diameter increment, height increment, and crown width increment. Parameters for all three dynamic expressions for growth were estimated using weighted, nonlinear three-stage least squares. This estimation method provided a predictive model with slight improvements in standard errors for two of the three equations (an average of 3% for height and diameter growth) and no improvement for crown width, when compared with two-stage least squares. The system includes competition from shrubs and hardwoods in predictions of height growth, diameter growth and crown width increment. This allows individual-tree/distance-independent simulator architecture to be extended to young plantations in southern Oregon and northern California.     

Direct impacts of sheep upon Douglas-fir trees in two agrosilvopastoral systems

Livestock may provide important service and production functions in agroforestry systems. However, use of livestock in conifer/improved pasture agrosilvopastoral systems is currently limited by concerns about potential damage to trees by livestock. Effects of sheep grazing on Douglas-fir (Pseudotsuga menziesii) trees in two patterns of sheep/pasture/conifer agroforest (cluster and grid plantations) were studied from 4 years after planting (1983) until the first precommercial thinning at age 10 years (1988). Trees averaged over 1 m in height when grazing began in summer 1983. Some browsing of tree lateral branches by sheep occurred regardless of grazing season in 1983–1985. However, the 2 to 10% of current year's lateral branch growth removed by grazing sheep was too low to impact tree growth. Sheep removed the terminal leaders from only 3 to 9% of trees each year during 1983–1985. Most browsing of terminals occurred in the summer when other forages had become mature and were relatively unpalatable to sheep. Less than 13% of agroforest trees were debarked by sheep each year during 1983–1987. By the end of grazing in 1987, less than 8% of agroforest trees had sustained a level of debarking likely to impact future growth (>50% of tree circumference debarked). Grazing had no discernible effect upon tree diameter or height in any year (P > 0.05). Total tree mortality attributable to sheep grazing during 1983–1987 was only 0.9%, including three trees girdled by sheep and two debarked trees which were subsequently attacked by insects. Overall, grazing had no detrimental impact on timber stand growth or mortality.Submitted as Oregon Agricultural Experiment Station Technical Paper No. 9628.     

Bole water content shows little seasonal variation in century-old Douglas-fir trees

Purportedly, large Douglas-fir trees in the American Pacific Northwest use water stored in bole tissues to ameliorate the effects of seasonal summer drought, the water content of bole tissues being drawn down over the summer months and replenished during the winter. Continuous monitoring of bole relative water content (RWC) in two 110-120-year-old Douglas-fir trees with ThetaProbe impedance devices provided an integrated measure of phloem-sapwood water content over 4 years. Seasonal changes in RWC closely tracked cambial activity and wood formation, but lagged changes in soil water content by 2-3 months. The RWC in the combined phloem and sapwood markedly increased during earlywood production in the late spring and early summer to maximum values of 64-77% as plant available soil water (ASW) decreased by approximately 60%. With transition and latewood formation, RWC decreased to minimum values of 59-72%, even as ASW increased in the fall. The difference between minimum RWC in the fall and maximum RWC in midsummer was only approximately 5%. Seasonal changes in bole RWC corresponded to cambial phenology, although decreasing AWS appeared to trigger the shift from earlywood to latewood formation.     

Respiratory potential in sapwood of old versus young ponderosa pine trees in the Pacific Northwest

Our primary objective was to present and test a new technique for in vitro estimation of respiration of cores taken from old trees to determine respiratory trends in sapwood. Our secondary objective was to quantify effects of tree age and stem position on respiratory potential (rate of CO2 production of woody tissue under standardized laboratory conditions). We extracted cores from one to four vertical positions in boles of +200-, +50- and +15-year-old Pinus ponderosa Dougl. ex Laws. trees. Cores were divided into five segments corresponding to radial depths of inner bark; outer, middle and inner sapwood; and heartwood. Data suggested that core segment CO2 production was an indicator of its respiratory activity, and that potential artifacts caused by wounding and extraction were minimal. On a dry mass basis, respiratory potential of inner bark was 3-15 times greater than that of sapwood at all heights for all ages (P < 0.0001). Within sapwood at all heights and in all ages of trees, outer sapwood had a 30-60% higher respiratory potential than middle or inner sapwood (P < 0.005). Heartwood had only 2-10% of the respiratory potential of outer sapwood. For all ages of trees, sapwood rings produced in the same calendar year released over 50% more CO2 at treetops than at bases (P < 0.0001). When scaled to the whole-tree level on a sapwood volume basis, sapwood of younger trees had higher respiratory potential than sapwood of older trees. In contrast, the trend was reversed when using the outer-bark surface area of stems as a basis for comparing respiratory potential. The differences observed in respiratory potential calculated on a core dry mass, sapwood volume, or outer-bark surface area basis clearly demonstrate that the resulting trends within and among trees are determined by the way in which the data are expressed. Although these data are based on core segments rather than in vivo measurements, we conclude that the relative differences are probably valid even if the absolute differences are not.     

Climate signal age effects—Evidence from young and old trees in the Swiss Engadin

A potential limitation of tree-ring based climate reconstructions is related to climate signal age effects (CSAE). CSAE may arise if the climatic response of young tree-rings differs from that of old tree-rings. This could mean that climatic signals become stronger (or weaker) with tree aging, or that the seasonality of signals or the sensitivity to a specific element (e.g., temperature, precipitation) changes over time. Such changes would affect the interpretation of dendroclimatic reconstructions, as the tree-rings included in these records are generally oldest at the end of a record (e.g., 21st century)—which is the time period generally used for calibration with instrumental data.We here addressed this concern by analyzing young and old Pinus cembra trees from three high elevation sites in the central European Alps. Core and disc samples were collected in pre-defined plots to allow for a representative analysis of tree ages with tree-ring width (TRW) measurement series categorized into age classes (i) >1880, (ii) 1880–1939, and (iii) 1940–2002. Notably we report on the signal of the very young category (iii) not yet described in literature, and thus allow estimation of climate response and signal strength characteristics during the first years of the trees’ lifespans.Comparison of age classes (i)–(iii) revealed differences in TRW coherence and size, but little change in climatic signal. CSAE are in the order of the differences recorded among high elevation sites—a conclusion that holds for inter-annual to decadal scale TRW variations at near-treeline Swiss stone pine. Such data are typically included in regional and larger-scale temperature reconstructions; thus, our results add confidence to long-term climate estimates integrating a range of tree-ring age classes. Other findings, such as the reaction wood in juvenile tree-rings, and sensitivity of the climate signal to sample replication, suggest that comparisons of young and old age classes, and separate calibration of these categories against instrumental climate data might further the estimation of long-term uncertainty changes in tree-ring based climate reconstructions.     

Belowground competition from overstory trees influences Douglas-fir sapling morphology in thinned stands

We evaluated effects of belowground competition on morphology of naturally established coast Douglas-fir (Pseudotsuga menziesii var. menziesii (Mirb.) Franco) saplings in 60- to 80-year-old thinned Douglas-fir stands in southwestern Washington. We separately quantified belowground competition from overstory and understory sources using trenching and understory removal. In this light-limited environment of 26 ± 16% (std. dev.) full sunlight, 2-year exclusion of tree root competition by trenching increased sapling stem biomass by 18%, total aboveground biomass by 21%, number of interwhorl buds by 68%, total foliar biomass by 33%, and foliar biomass on branch components over 4 years old by 143%. Belowground competition did not influence shoot:root ratio or foliar efficiency (i.e., stem growth per unit foliage biomass). Sapling needle size, specific leaf area, and internodal distance also were not affected by belowground competition; these variables were apparently a function of the low-light environment. The principal source of belowground competition was roots of overstory trees; effects of belowground competition from understory vegetation were minor. Thus, under a partial overstory, morphology of Douglas-fir regeneration was influenced by both belowground and aboveground competition from overstory trees. In this environment, understory vegetation control would not likely influence belowground competition to an extent that would affect sapling morphology.     

Branch mortality and potential litterfall from Douglas-fir trees in stands of varying density

Forest floor characteristics influence nutrient cycling and energy flow properties of forest ecosystems, and determine quality of habitat for many forest plants and animals. Differential crown recession and crown development among stands of differing density suggest that an opportunity may exist to control the input of fine woody litter into the system by manipulating stand density. The objective was to measure the rate of branch mortality among stands of differing density and to estimate the range in total per hectare necromass inputs. Although litter traps are reliable for estimating per hectare rates of litterfall, branch mortality dating on sectioned stems uniquely allows assessment of several other litterfall components: (1) individual tree contributions to total litterfall; (2) the amount of branch material released by mortality, regardless of whether the branches are shed to the forest floor; (3) the distribution of basal diameters characterizing the litterfall from a given tree and stand. Twenty-four trees were felled and sectioned on permanent plots that were part of a silvicultural study of stand density regimes in Douglas-fir (Pseudotsuga menziesii (Mirbel) Franco.). Whorl branches were dissected out of bole sections to determine the dates of mortality, and a branch biomass equation was applied to estimate potential rate of litterfall. Periodic annual rates were expressed in four ways: (1) number of branches per tree; (2) mass of branches per tree; (3) mass of branches per unit of crown projection area; (4) mass of branches per hectare. For the growth periods investigated, larger trees and trees growing on denser plots tended to release a greater necromass through branch mortality. Average branch basal diameter generally decreased with increasing stand density. Annual branch mortality ranged from 33 to 430 g m⁻² crown projection area for individual trees, and from 236 to 1035 kg ha⁻¹ for individual plots. These rates approached the low end of the range of previously published fine litterfall rates for Douglas-fir. Rates on these plots were relatively low owing to the temporary delay in crown recession imposed by artificial thinning. A conceptual model of branch litter dynamics is presented to depict consistencies with crown development among stands managed under different density regimes.     

The geographic origin of old Douglas-fir stands growing in Central Europe

Since the nineteenth century, Douglas-fir seed sources have been widely used for establishment of forest stands outside its natural distribution range. In Europe, some of these old Douglas-fir stands are registered as seed stands and provide seed sources for nurseries, although it is unclear from which region in North America they originate. In recent years, the interest in planting Douglas-fir has increased substantially because the species is seen as a potential adaptation option to climate change. This makes the assignment of European Douglas-fir stands of unknown seed origin to their geographic origin in North America increasingly important, because the genetic quality of these plantations must be guaranteed. In this study, we use 13 nuSSR loci to investigate the origin of 67 Austrian and German Douglas-fir stands of unknown origin. We performed a hierarchical Bayesian cluster analysis using 38 native Douglas-fir populations. The resulting clusters are used as reference populations to assign the 67 Central European Douglas-fir stands from Austria and Germany planted more than 80 years ago. Our results suggest that the majority of our investigated Douglas-fir stands come from central Washington (USA), the recommended seed zones for Central Europe. Some stands were located outside the suggested area, e.g. central Oregon and Santa Fe (New Mexico). The accuracy assessment of our approach revealed the best performance for the highest hierarchical level, e.g. assigning populations either to the coastal or the Rocky Mountain variety. As expected, the uncertainty increases with decreasing hierarchical level. The final assessment, if an admixture of seed sources within the European Douglas-fir stands is evident suggests that 23 of the Douglas-fir stands show an admixture which was not detected in our Douglas-fir reference populations growing in the natural distribution range.     

Correlations between stable carbon-isotope abundance and hydraulic conductivity in Douglas-fir across a climate gradient in Oregon, USA

Stomatal conductance in trees is related to both foliar carbon-isotope abundance and stem hydraulic properties. By combining these relationships, I hypothesized that carbon-isotope abundance in foliage should vary with limitations to water movement through supporting branches. I sampled Douglas-fir branches (Pseudotsuga menziesii (Mirb.) Franco) from six sites across a climate gradient in Oregon, USA for foliar carbon-isotope abundance and stem hydraulic properties. I used a forest growth model to quantify climate-induced stomatal limitations, expressed as reduced potential transpiration, across the gradient. Foliar stable carbon-isotope abundance showed a strong inverse relationship with branch specific conductivity (hydraulic conductivity per unit functional sapwood area) and leaf-specific conductivity (hydraulic conductivity per unit leaf area). Foliar stable carbon-isotope abundance was correlated with modeled reductions in potential transpiration; however, the inclusion of leaf-specific conductivity improved the correlation by more than 30%. Combined, leaf-specific conductivity and climate-induced stomatal constraints explained 84% of the variation in foliar isotope abundance in 1994 foliage. This model was confirmed on foliage classes 1990-1993.