Converging patterns of uptake and hydraulic redistribution of soil water in contrasting woody vegetation types

We used concurrent measurements of soil water content and soil water potential (Psi(soil)) to assess the effects of Psi(soil) on uptake and hydraulic redistribution (HR) of soil water by roots during seasonal drought cycles at six sites characterized by differences in the types and amounts of woody vegetation and in climate. The six sites included a semi-arid old-growth ponderosa pine (Pinus ponderosa Dougl. ex P. Laws & C. Laws) forest, a moist old-growth Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) forest, a 24-year-old Douglas-fir forest and three Brazilian savanna sites differing in tree density. At all of the sites, HR was confined largely to the upper 60 cm of soil. There was a common threshold relationship between the relative magnitude of HR and Psi(soil) among the six study sites. Below a threshold Psi(soil) of about -0.4 MPa, overnight recharge of soil water storage increased sharply, and reached a maximum value of 80-90% over a range of Psi(soil) from ~ -1.2 to -1.5 MPa. Although amounts of water hydraulically redistributed to the upper 60 cm of soil were relatively small (0 to 0.4 mm day(-1)), they greatly reduced the rates of seasonal decline in Psi(soil). The effectiveness of HR in delaying soil drying diminished with increasing sapwood area per ground area. The relationship between soil water utilization and Psi(soil) in the 20-60-cm layer was nearly identical for all six sites. Soil water utilization varied with a surrogate measure of rhizosphere conductance in a similar manner at all six sites. The similarities in relationships between Psi(soil) and HR, soil water utilization and relative rhizosphere conductance among the six sites, suggests that, despite probable differences in maximum rooting depth and density, there was a convergence in biophysical controls on soil water utilization and redistribution in the upper soil layers where the density of finer roots is greatest.     

Effects of branch height on leaf gas exchange,branch hydraulic conductance and branch sap flux in open-grown ponderosa pine

Recent studies have shown that stomata respond to changes in hydraulic conductance of the flow path from soil to leaf. In open-grown tall trees, branches of different heights may have different hydraulic conductances because of differences in path length and growth. We determined if leaf gas exchange, branch sap flux, leaf specific hydraulic conductance, foliar carbon isotope composition (delta13C) and ratios of leaf area to sapwood area within branches were dependent on branch height (10 and 25 m) within the crowns of four open-grown ponderosa pine (Pinus ponderosa Laws.) trees. We found no difference in leaf gas exchange or leaf specific hydraulic conductance from soil to leaf between the upper and lower canopy of our study trees. Branch sap flux per unit leaf area and per unit sapwood area did not differ between the 10- and 25-m canopy positions; however, branch sap flux per unit sapwood area at the 25-m position had consistently lower values. Branches at the 25-m canopy position had lower leaf to sapwood area ratios (0.17 m2 cm-2) compared with branches at the 10-m position (0.27 m2 cm-2) (P = 0.03). Leaf specific conductance of branches in the upper crown did not differ from that in the lower crown. Other studies at our site indicate lower hydraulic conductance, sap flux, whole-tree canopy conductance and photosynthesis in old trees compared with young trees. This study suggests that height alone may not explain these differences.     

Water limitations to carbon exchange in old-growth and young ponderosa pine stands

We investigated the impact of seasonal soil water deficit on the processes driving net ecosystem exchange of carbon (NEE) in old-growth and recently regenerating ponderosa pine (Pinus ponderosa Doug. ex Laws.) stands in Oregon. We measured seasonal patterns of transpiration, canopy conductance and NEE, as well as soil water, soil temperature and soil respiration. The old-growth stand (O) included two primary age classes (50 and 250 years), had a leaf area index (LAI) of 2.1 and had never been logged. The recently regenerating stand (Y) consisted predominantly of 14-year-old ponderosa pine with an LAI of 1.0. Both stands experienced similar meteorological conditions with moderately cold wet winters and hot dry summers. By August, soil volumetric water content within the upper 30 cm had declined to a seasonal minimum of 0.07 at both sites. Between April and June, both stands showed similar rates of transpiration peaking at 0.96 mm day(-1); thereafter, trees at the Y site showed increasing drought stress with canopy stomatal resistance increasing 6-fold by mid-August relative to values for trees at the O site. Over the same period, predawn water potential (psi(pd)) of trees at the Y site declined from -0.54 to -1.24 MPa, whereas psi(pd) of trees at the O site remained greater than -0.8 MPa throughout the season. Soil respiration at the O site showed a strong seasonal correlation with soil temperature with no discernible constraints imposed by declining soil water. In contrast, soil respiration at the Y site peaked before seasonal maximal soil temperatures and declined thereafter with declining soil water. No pronounced seasonal pattern in daytime NEE was observed at either site between April and September. At the Y site this behavior was driven by concurrent soil water limitations on soil respiration and assimilation, whereas there was no evidence of seasonal soil water limitations on either process at the O site.     

Predicting the decline in daily maximum transpiration rate of two pine stands during drought based on constant minimum leaf water potential and plant hydraulic conductance

The effect of drought on forest water use is often estimated with models, but comprehensive models require many parameters, and simple models may not be sufficiently flexible. Many tree species, Pinus species in particular, have been shown to maintain a constant minimum leaf water potential above the critical threshold for xylem embolism during drought. In such cases, prediction of the relative decline in daily maximum transpiration rate with decreasing soil water content is relatively straightforward. We constructed a soil-plant water flow model assuming constant plant conductance and daily minimum leaf water potential, but variable conductance from soil to root. We tested this model against independent data from two sites: automatic shoot chamber data and sap flow measurements from a boreal Scots pine (Pinus sylvestris L.) stand; and sap flow measurements from a maritime pine (Pinus pinaster Ait.) stand. To focus on soil limitations to water uptake, we expressed daily maximum transpiration rate relative to the rate that would be obtained in wet soil with similar environmental variables. The comparison was successful, although the maritime pine stand showed carry-over effects of the drought that we could not explain. For the boreal Scots pine stand, daily maximum transpiration was best predicted by water content of soil deeper than 5 cm. A sensitivity analysis revealed that model predictions were relatively insensitive to the minimum leaf water potential, which can be accounted for by the importance of soil resistance of drying soil. We conclude that a model with constant plant conductance and minimum leaf water potential can accurately predict the decline in daily maximum transpiration rate during drought for these two pine stands, and that including further detail about plant compartments would add little predictive power, except in predicting recovery from severe drought.     

Restoration thinning and influence of tree size and leaf area to sapwood area ratio on water relations of Pinus ponderosa

Ponderosa pine (Pinus ponderosa Dougl. ex P. Laws) forest stand density has increased significantly over the last century (Covington et al. 1997). To understand the effect of increased intraspecific competition, tree size (height and diameter at breast height (DBH)) and leaf area to sapwood area ratio (A(L):A(S)) on water relations, we compared hydraulic conductance from soil to leaf (kl) and transpiration per unit leaf area (Q(L)) of ponderosa pine trees in an unthinned plot to trees in a thinned plot in the first and second years after thinning in a dense Arizona forest. We calculated kl and Q(L) based on whole- tree sap flux measured with heat dissipation sensors. Thinning increased tree predawn water potential within two weeks of treatment. Effects of thinning on kl and Q(L) depended on DBH, A(L):A(S) and drought severity. During severe drought in the first growing season after thinning, kl and Q(L) of trees with low A(L):A(S) (160-250 mm DBH; 9-11 m height) were lower in the thinned plot than the unthinned plot, suggesting a reduction in stomatal conductance (g(s)) or reduced sapwood specific conductivity (K(S)), or both, in response to thinning. In contrast kl and Q(L) were similar in the thinned plot and unthinned plot for trees with high A(L):A(S) (260-360 mm DBH; 13-16 m height). During non-drought periods, kl and Q(L) were greater in the thinned plot than in the unthinned plot for all but the largest trees. Contrary to previous studies of ponderosa pine, A(L):A(S) was positively correlated with tree height and DBH. Furthermore, kl and Q(L) showed a weak negative correlation with tree height and a strong negative correlation with A(S) and thus A(L):A(S) in both the thinned and unthinned plots, suggesting that trees with high A(L):A(S) had lower g(s). Our results highlight the important influence of stand competitive environment on tree-size-related variation in A(L):A(S) and the roles of A(L):A(S) and drought on whole-tree water relations in response to thinning.     

Xylem vulnerability to cavitation in Pseudotsuga menziesii and Pinus ponderosa from contrasting habitats

In the Rocky Mountains, ponderosa pine (Pinus ponderosa (ssp.) ponderosa Dougl. ex P. Laws. & C. Laws) often co-occurs with Douglas-fir (Pseudotsuga menziesii var. glauca (Mayr) Franco). Despite previous reports showing higher shoot vulnerability to water-stress-induced cavitation in ponderosa pine, this species extends into drier habitats than Douglas-fir. We examined: (1) whether roots and shoots of ponderosa pine in riparian and slope habitats are more vulnerable to water-stress-induced cavitation than those of Douglas-fir; (2) whether species-specific differences in vulnerability translate into differences in specific conductivity in the field; and (3) whether the ability of ponderosa pine to extend into drier sites is a result of (a) greater plasticity in hydraulic properties or (b) functional or structural adjustments. Roots and shoots of ponderosa pine were significantly more vulnerable to water-stress-induced cavitation (overall mean cavitation pressure, Psi(50%) +/- SE = -3.11 +/- 0.32 MPa for shoots and -0.99 +/- 0.16 MPa for roots) than those of Douglas-fir (Psi(50%) +/- SE = -4.83 +/- 0.40 MPa for shoots and -2.12 +/- 0.35 MPa for roots). However, shoot specific conductivity did not differ between species in the field. For both species, roots were more vulnerable to cavitation than shoots. Overall, changes in vulnerability from riparian to slope habitats were small for both species. Greater declines in stomatal conductance as the summer proceeded, combined with higher allocation to sapwood and greater sapwood water storage, appeared to contribute to the ability of ponderosa pine to thrive in dry habitats despite relatively high vulnerability to water-stress-induced cavitation.     

Container seedlings outperform barefoot stock: Survival and growth after 10 years

Ponderosa pine, Jeffrey pine, and Douglas-fir seedlings were planted in container or bareroot form at three elevations in northern California. At the lowest elevation (762 m), container seedlings of ponderosa pine were significantly taller than bareroot seedlings at ages 4 and 10, and had breast-height diameters that were significantly larger than bareroot counterparts at age 10. Survival of Douglas-fir container seedlings was significantly greater than that of barefoot seedlings for all ages tested. At the mid-elevation site (1220 m), container seedlings of ponderosa pine and Douglas-fir were significantly larger in breast-height diameter than bareroot seedlings at age 10. Douglas-fir container seedlings survived significantly better at all ages than barefoot seedlings. At the highest elevation (1662 m), seedling height and diameter did not differ significantly, but survival of container seedlings was significantly higher than barefoot seedlings for both pine species at all ages tested.     

Reliance on stored water increases with tree size in three species in the Pacific Northwest

In tall old forests, limitations to water transport may limit maximum tree height and reduce photosynthesis and carbon sequestration. We evaluated the degree to which tall trees could potentially compensate for hydraulic limitations to water transport by increased use of water stored in xylem. Using sap flux measurements in three tree species of the Pacific Northwest, we showed that reliance on stored water increases with tree size and estimated that use of stored water increases photosynthesis. For Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), water stored in xylem accounted for 20 to 25% of total daily water use in 60-m trees, whereas stored water comprised 7% of daily water use in 15-m trees. For Oregon white oak (Quercus garryana Dougl. ex Hook.), water stored in xylem accounted for 10 to 23% of total daily water use in 25-m trees, whereas stored water comprised 9 to 13% of daily water use in 10-m trees. For ponderosa pine (Pinus ponderosa Dougl. ex Laws.), water stored in xylem accounted for 4 to 20% of total daily water use in 36-m trees, whereas stored water comprised 2 to 4% of daily water use in 12-m trees. In 60-m Douglas-fir trees, we estimated that use of stored water supported 18% more photosynthesis on a daily basis than would occur if no stored water were used, whereas 15-m Douglas-fir trees gained 10% greater daily photosynthesis from use of stored water. We conclude that water storage plays a significant role in the water and carbon economy of tall trees and old forests.     

Carbon dioxide and water vapor exchange by young and old ponderosa pine ecosystems during a dry summer

We investigated key factors controlling mass and energy exchange by a young (6-year-old) ponderosa pine (Pinus ponderosa Laws.) plantation on the west side of the Sierra Nevada Mountains and an old-growth ponderosa pine forest (mix of 45- and 250-year-old trees) on the east side of the Cascade Mountains, from June through September 1997. At both sites, we operated eddy covariance systems above the canopy to measure net ecosystem exchange of carbon dioxide and water vapor, and made concurrent meteorological and ecophysiological measurements. Our objective was to understand and compare the controls on ecosystem processes in these two forests. Precipitation is much higher in the young plantation than in the old-growth forest (1660 versus 550 mm year-1), although both forests experienced decreasing soil water availability and increasing vapor pressure deficits (D) as the summer of 1997 progressed. As a result, drought stress increased at both sites during this period, and changes in D strongly influenced ecosystem conductance and net carbon uptake. Ecosystem conductance for a given D was higher in the young pine plantation than in the old-growth forest, but decreased dramatically following several days of high D in late summer, possibly because of xylem cavitation. Net CO2 exchange generally decreased with conductance at both sites, although values were roughly twice as high at the young site. Simulations with the 3-PG model, which included the effect of tree age on fluxes, suggest that, during the fall through spring period, milder temperatures and ample water availability at the young site provide better conditions for photosynthesis than at the old pine site. Thus, over the long-term, the young site can carry more leaf area, and the climatic conditions between fall and spring offset the more severe limitations imposed by summer drought.     

Hydraulic redistribution of soil water by neotropical savanna trees

The magnitude and direction of water transport by the roots of eight dominant Brazilian savanna (Cerrado) woody species were determined with a heat pulse system that allowed bidirectional measurements of sap flow. The patterns of sap flow observed during the dry season in species with dimorphic root systems were consistent with the occurrence of hydraulic redistribution of soil water, the movement of water from moist to drier regions of the soil profile via plant roots. In these species, shallow roots exhibited positive sap flow (from the soil into the plant) during the day and negative sap flow (from the plant into the soil) during the night. Sap flow in the taproots was positive throughout the 24-h period. Diel fluctuations in soil water potential, with maximum values occurring at night, provided evidence for partial rewetting of upper soil layers by water released from shallow roots. In other species, shallow roots exhibited negative sap flow during both the day and night, indicating that hydraulic redistribution was occurring continuously. A third sap flow pattern was observed at the end of the dry season after a heavy rainfall event when sap flow became negative in the taproot, and positive in the small roots, indicating movement of water from upper soil layers into shallow roots, and then into taproots and deeper soil layers. Experimental manipulations employed to evaluate the response of hydraulic redistribution to changes in plant and environmental conditions included watering the soil surface above shallow roots, decreasing transpiration by covering the plant and cutting roots where probes were inserted. Natural and manipulated patterns of sap flow in roots and stems were consistent with passive movement of water toward competing sinks in the soil and plant. Because dry shallow soil layers were often a stronger sink than the shoot, we suggest that the presence of a dimorphic root system in deciduous species may play a role in facilitating leaf expansion near the end of the dry season when the soil surrounding shallow lateral roots is still dry.     

Regulation of ponderosa pine foliar physiology and insect resistance mechanisms by basal area treatments

We compared foliar physiology and several measures of tree resistance to insect attack among ponderosa pine (Pinus ponderosa var. scopulorum Engelm.) trees growing in thinned stands. Measurements were made in a second-growth ponderosa pine forest in northern Arizona where the basal area treatments (6.9, 18.4, 27.6, 78.2 m(2) ha(-1)) have been experimentally maintained by frequent thinnings for 32 years before our measurements began in 1994. Most of the physiological characteristics measured were affected by the basal area treatments. As stand basal area increased from 6.9 to 78.2 m(2) ha(-1), predawn water potential, midday water potential, net photosynthetic rate, resin production, phloem thickness, and foliar toughness decreased. Foliar nitrogen concentration was greatest in trees in the intermediate basal area treatments. Our results show that the physiological condition of second-growth ponderosa pine can be manipulated by silvicultural control of stand basal area, and support the hypothesis that high stand basal area increases tree stress and decreases tree resistance to insect attack.     

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.     

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.     

Co-occurring species differ in tree-ring delta(18)O trends

The stable oxygen isotope ratio (delta(18)O) of tree-ring cellulose is jointly determined by the delta(18)O of xylem water, the delta(18)O of atmospheric water vapor, the humidity of the atmosphere and perhaps by species-specific differences in leaf structure and function. Atmospheric humidity and the delta(18)O of water vapor vary seasonally and annually, but if the canopy atmosphere is well mixed, atmospheric characteristics should be uniform among co-occurring trees. In contrast, xylem water delta(18)O is determined by the delta(18)O of water being drawn from the soil, which varies with depth. If co-occurring trees draw water from different soil depths, this soil-water delta(18)O signal would be manifest as differences in delta(18)O among the trees. We examined the variation in tree ring delta(18)O, over eight decades during the 20th Century, among three species co-occurring in natural forest stands of the northern Rocky Mountains in the USA. We sampled 10 Douglas-firs (Pseudotsuga menziesii (Mirb.) Franco var. glauca), 10 ponderosa pines (Pinus ponderosa Laws.) and seven western white pines (Pinus monticola Dougl.). As expected, variation in atmospheric conditions was recorded in the delta(18)O of the cellulose produced in a given year, but observed climatic correlations with delta(18)O were weak. Significant correlations with June climate data included: daily maximum temperature (r = 0.29), daily minimum temperature (r = -0.25), mean temperature (r = 0.20), mean daily precipitation (r = -0.54), vapor pressure deficit (r = 0.32) and solar radiation (r = 0.44). Lagged effects were observed in Douglas-fir and western white pine. In these species, the delta(18)O of a given annual ring was correlated with the delta(18)O of the previous ring. Ponderosa pine showed no significant autocorrelation. Although the species means were correlated among years (r = 0.67 to 0.76), ponderosa pine was consistently enriched in delta(18)O relative to the other species; differences were close to 2 per thousand and they are steadily increasing. Relative to the mean for the three species, ponderosa pine is becoming steadily more enriched (-1.0 per thousand). In contrast, Douglas-fir is being steadily depleted and western pine is intermediate, with an enrichment of 0.5 per thousand. Because all trees were exposed to the same atmospheric conditions, the differences in delta(18)O observed between species are likely due either to differences in the depth of water extraction or leaf function. If the former, presumably ponderosa pine has steadily taken up more water from near the soil surface and Douglas-fir has shifted uptake to a greater depth. If the latter, we suggest the pronounced changes in leaf-water delta(18)O are a result of changes in leaf structure and function with tree size and age.     

Truffle abundance and mycophagy by northern flying squirrels in eastern Washington forests

Although much is known about truffle abundance and rodent mycophagy in mesic Douglas-fir forests in the Pacific Northwest, few data are available for dry interior montane forests dominated by ponderosa pine (Pinus ponderosa), Douglas-fir (Pseudotsuga menziesii), and grand fir (Abies grandis). Our objective was to quantify the relationship between the abundance and diversity of ectomycorrhizal fungal sporocarps in the soil and in the diets of northern flying squirrels (Glaucomys sabrinus) in low-elevation forests of the eastern Washington Cascades. We randomly sampled four stands each of three cover types: dry open ponderosa pine, mesic young mixed-conifer forest, and mesic mature mixed-conifer forest. We sampled the soil for hypogeous sporocarps during the spring of 1999 and 2000. We collected fecal pellets from 318 flying squirrels live-trapped during the fall of 1997–2000. We sampled 2400 m² of soil surface and found truffles in 40% of 600 plots. Total biomass collected was 609 g. Spring truffle biomass on a kg/ha basis averaged 1.72 in open pine, 3.56 in young, and 4.11 in mature forest. Twenty-two species were collected across all cover types, with all but three species belonging to the Basidiomycotina. Eleven dominant species accounted for 91–94% of truffle biomass in each cover type. Four dominant species accounted for 60–70% of spring truffle biomass: Gautieria monticola, Hysterangium coriaceum, Rhizopogon parksii, and R. vinicolor. Truffle assemblages, richness and total biomass differed among cover types: richness and biomass were highest in young and mature mixed-conifer forest, and lowest in open ponderosa pine forest. Fall squirrel diets were composed of 23 genera or groups of fungi, plus about 22% plant material. Rhizopogon was the most abundant genus in the diet, followed by plant material, then Gautieria, Leucogaster, Alpova, and Hysterangium. Diets in different cover types were similar in the composition, richness, evenness, and the ratio of fungus to plant material. Diet richness varied over the study period. Nineteen truffle genera were detected in fall fecal samples versus 12 in spring soil samples. Management of low-elevation dry forest to maintain or restore stable fire regimes might reduce truffle diversity at stand scales by simplifying stand composition and structure; but, such management might increase long-term beta and landscape truffle diversity and persistence by reducing the occurrence of high-intensity fires and stabilizing inherent fire disturbance regimes.     

Effect of soil temperature on root and shoot growth and on budburst timing in conifer seedling transplants

Root growth in seedling transplants of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), Pacific silver fir (Abies amabilis(Dougl.) Forbes), noble fir (Abies procera Rehd.), lodgepole pine (Pinus contorta Dougl. ex Loud.) and ponderosa pine (Pines ponderosa Doug. ex Laws.) began when soil temperature exceeded 5 °C. Root growth increased rapidly after 10 °C and attained maximum values at 20 °C. At 30 °C, no root growth occurred in the firs; in the pines, root growth was 30 to 39% of maximum. Maximum shoot growth also occurred at 20 °C. In ponderosa pine, height growth of seedlings from a high-elevation source was unaffected by cold soil, but in low-elevation seedlings it was reduced. Budburst in Douglas-fir and the pines was delayed up to 11 days by cold soil, whereas in silver fir and noble fir, it was only slightly delayed. Prior to new root growth in ponderosa pine, xylem pressure potentials and stomatal conductances during the afternoon indicated reduced stomatal opening at all soil temperatures, whereas 23 days later, stomata were open to a greater degree when temperatures exceeded 10 °C.     

The fate of hydraulically redistributed water in a semi-arid zone eucalyptus species

Although hydraulic redistribution has been observed for a range of tree species, including Eucalyptus kochii subsp. borealis (C. Gardner) D. Nicolle, there is limited direct evidence that water taken up by deep roots in moist soil is in fact exuded by shallow roots in dry soil. This paper reports an experiment designed to test this hypothesis. Water enriched with deuterium was added to the groundwater via a slotted tube at 4.5 m depth below 5-year-old E. kochii subsp. borealis trees. Nocturnal sap flow increased markedly immediately after deep irrigation, indicating that the trees were using water from this depth. Two weeks later, samples of surface soil and xylem water were found to contain levels of deuterium up to 30% higher than soils and xylem water from a control plot upslope of the main treatment plot. This is strong evidence that trees used groundwater and that efflux of important amounts of hydraulically redistributed water occurred via the roots of E. kochii subsp. borealis.     

Age-related changes in ecosystem structure and function and effects on water and carbon exchange in ponderosa pine

As forests age, their structure and productivity change, yet in some cases, annual rates of water loss remain unchanged. To identify mechanisms that might explain such observations, and to determine if widely different age classes of forests differ functionally, we examined young (Y, approximately 25 years), mature (M, approximately 90 years) and old (O, approximately 250 years) ponderosa pine (Pinus ponderosa Dougl. ex P. Laws.) stands growing in a drought-prone region of central Oregon. Although the stands differed in tree leaf area index (LAIT) (Y = 0.9, M = 2.8, O = 2.1), cumulative tree transpiration measured by sap flow did not differ substantially during the growing season (100-112 mm). Yet when water was readily available, transpiration per unit leaf area of the youngest trees was about three times that of M trees and five times that of O trees. These patterns resulted from a nearly sixfold difference in leaf specific conductance (KL) between the youngest and oldest trees. At the time of maximum transpiration in the Y stand in May-June, gross carbon uptake (gross ecosystem production, GEP) was similar for Y and O stands despite an almost twofold difference in stand leaf area index (LAIS). However, the higher rate of water use by Y trees was not sustainable in the drought-prone environment, and between spring and late summer, KL of Y trees declined fivefold compared with a nearly twofold decline for M trees and a < 30% reduction in O trees. Because the Y stand contained a significant shrub understory and more exposed soil, there was no appreciable difference in mean daily latent energy fluxes between the Y stand and the older stands as measured by the eddy-covariance technique. These patterns resulted in 60 to 85% higher seasonal GEP and 55 to 65% higher water-use efficiency at the M and O stands compared with the Y stand.     

Modeling daily gas exchange of a Douglas-fir forest: comparison of three stomatal conductance models with and without a soil water stress function

Modeling stomatal conductance is a key element in predicting tree growth and water use at the stand scale. We compared three commonly used models of stomatal conductance, the Jarvis-Loustau, Ball-Berry and Leuning models, for their suitability for incorporating soil water stress into their formulation, and for their performance in modeling forest ecosystem fluxes. We optimized the parameters of each of the three models with sap flow and soil water content data. The optimized Ball-Berry model showed clear relationships with air temperature and soil water content, whereas the optimized Leuning and Jarvis-Loustau models only showed a relationship with soil water content. We conclude that use of relative humidity instead of vapor pressure deficit, as in the Ball-Berry model, is not suitable for modeling daily gas exchange in Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) in the Speulderbos forest near the village of Garderen, The Netherlands. Based on the calculated responses to soil water content, we linked a model of forest growth, FORGRO, with a model of soil water, SWIF, to obtain a forest water-balance model that satisfactorily simulated carbon and water (transpiration) fluxes and soil water contents in the Douglas-fir forest for 1995.     

Evaluation of transpiration in a Douglas-fir stand by means of sap flow measurements

Transpiration of a Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) stand was evaluated by sap flow measurements during a 4-month period. Between-tree variation in sap flow depended on crown class. On a sunny day, total transpiration was 1.6, 8.0 and 22.0 liters day(-1) for suppressed, codominant and dominant trees, respectively. Transpiration estimated by sap flow fell below potential evapotranspiration when available soil water decreased below 30% of its maximum value. Sap flow measurements gave transpiration values similar to those obtained by the water balance method.