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.

Effects of light on shoot geometry and needle morphology in Abies amabilis.

In some conifers, shoot geometry and needle morphology vary significantly in response to the light conditions under which they develop. We measured shoot length, silhouette area, total projected needle area, total needle weight and needle thickness on current shoots developed under a wide range of light conditions in a 36-year-old Abies amabilis (Dougl.) Forbes stand. Current light was quantified by evaluating percent openness from hemispherical photographs taken before the growing season. Unweighted total openness was correlated with shoot geometry and needle morphology better than any weighted indices tested. Needle thickness and leaf mass/area were both closely correlated with total openness (R(2) = 0.86 and 0.82, respectively). The most exposed needles were 2.5 times thicker and had 3-4 times more leaf mass/area than the most shaded needles. Total projected leaf area/shoot silhouette area was also correlated with openness (R(2) = 0.74) and was about twice as high in sun shoots as in shaded shoots. As a result of greater leaf mass/leaf area and greater leaf area/shoot silhouette area, a unit of intercepted light was dispersed over about 6 times as much leaf mass in a sun shoot as in a shade shoot, which presumably permits more efficient utilization of the intercepted light under high illumination with less energy wastage to light saturation. Moreover, leaf mass per unit of silhouette area was almost exactly proportional to canopy openness, as predicted by resource optimization theory if nitrogen concentration and photosynthetic capacity per unit mass are constant in new leaves. The close correlation of needle thickness and leaf mass/area with openness suggests that either parameter could be used as an index of the distribution of light or light-driven processes in an A. amabilis canopy.

A comparison of measured and modeled ozone uptake into plant leaves.

Ozone uptake into plant leaves was measured in gas exchange chambers using a mass balance and a variable conductance approach. The variable conductance approach was found to more reliably measure ozone flux through stomata. Measurements using this approach were contrasted with estimates obtained by measuring stomatal conductance g(sw) and modeling ozone uptake using a diffusion equation, assuming a negligible ozone concentration in the substomatal cavity. Actual measurements of uptake were close, but slightly higher than modeled values, providing some support to the idea that substomatal ozone concentrations are close to zero. However, the difference between measured and modeled uptake values suggests either that (i) variable conductance approach measures more ozone uptake than caused by stomatal uptake alone or (ii) ozone conductance is underestimated.

Water flux in a hybrid poplar stand.

We studied water flux in a four-year-old stand of hybrid Populus during midsummer 1992. Study trees ranged in height from 11.0 to 15.1 m and in diameter from 8.3 to 15.1 cm. The large-leafed Populus hybrid was relatively poorly coupled to the atmosphere. The average value of the stomatal decoupling coefficient, Omega, was 0.66, indicating that, on average, a 10% change in stomatal conductance would result in only a 3 to 4% change in transpiration. During the middle of the summer, the smallest study tree used between 20 and 26 kg of water per day, whereas the largest tree used between 39 and 51 kg day(-1). The maximum observed rate of stand water loss was 4.8 mm day(-1) in this Populus clone. Maximum rates of sap velocity within the xylem were as high as 12.5 m h(-1); measured rates for exposed sunlit branches approached 90% of this maximum. Within-canopy patterns of stomatal conductance generally reflected patterns of incident radiation. Stomatal conductance of foliage grown in shade, even when exposed to non-limiting light and water source conditions, did not increase appreciably. Patterns of stomatal conductance under limiting and non-limiting conditions suggested that both stomatal conductance and leaf specific hydraulic conductivity (LSHC) were linked with the ability to exploit the light resource.

Acclimation responses of mature Abies amabilis sun foliage to shading.

This paper addresses two main questions. First, can evergreen foliage that has been structurally determined as sun foliage acclimate physiologically when it is shaded? Second, is this acclimation independent of the foliage ageing process and source-sink relations? To investigate these questions, a shading and debudding experiment was established using paired branches on opengrown Abies amabilis trees. For each tree, one branch was either shaded, debudded, or both, from before budbreak until the end of summer, while the other branch functioned as a control. Foliage samples were measured both prior to and during treatment for photosynthesis at light saturation (A max), dark respiration, nitrogen content, chlorophyll content, chlorophyll-to-nitrogen ratio and chlorophyll a:b ratio. All age classes of foliage responded similarly during the treatment, although pre-treatment values differed between age classes. Within 1 month after the treatment began, A max was lower in shaded foliage and remained lower throughout the treatment period. For debudded branches, A max was lower than the controls only during active shoot elongation. At the end of the treatments in September, A max in shade-treated sun foliage matched the rates in the true shade-formed foliage, but nitrogen remained significantly higher. By 1.5 months after treatment, chlorophyll content in shaded foliage was higher than in controls, and the chlorophyll a:b ratio was lower for the shaded foliage. On debudded branches, chlorophyll content and chlorophyll a:b ratio were similar to the values in control samples. Shading lowered the rate of nitrogen accumulation within a branch, while removing debudding decreased the amount of sequestered N that was exported from the older foliage to supply new growth. By September, chlorophyll content in shade-treated foliage was higher than that in the control sun foliage or in true shade foliage. The chlorophyll increase as a result of shading was unexpected. However, the chlorophyll-to-nitrogen ratio was identical for the shade-treated sun foliage and the true shade foliage while being significantly lower than the control sun foliage. It appears that acclimation to shading in mature foliage involves a reallocation of nitrogen within the leaf into thylakoid proteins. A redistribution of resources (nitrogen) among leaves is secondary and appears to function on a slower time scale than reallocation within the leaf. Thus, A. amabilis foliage that is structurally determined as sun foliage can acclimate to shade within a few months; this process is most likely independent of ageing and is only slightly affected by source-sink relations within a branch.

Effects of foliar nitrogen concentration on photosynthesis and water use efficiency in Douglas-fir.

Leaf-level physiological processes were studied in Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) to determine whether apparent increases in stand-level water use efficiency (WUE) observed in response to nitrogen (N) fertilization were attributable to foliar N effects on carbon fixation rates or on stomatal control of water loss. Photosynthesis and transpiration were measured at different light intensities and ambient CO(2) molar fractions and comparisons were made between current-year shoots with average foliar N concentrations of 1.58% (High-N) and 1.25% (Low-N). Photosynthetic rates and foliar N concentrations were positively correlated. In response to light, photosynthesis and stomatal conductance were closely coupled and a similar coupling was observed in response to different ambient CO(2) concentrations. Partitioning the photosynthetic responses into mesophyll and stomatal components indicated that foliar N altered mesophyll conductance but not stomatal control of water loss. High-N shoots had significantly greater rates of photosynthesis and transpiration than Low-N shoots and, as a result, instantaneous WUE did not differ significantly between High-N and Low-N shoots.

Influence of soil water on the physiological and morphological components of plant water balance in Populus trichocarpa, Populus deltoides and their F(1) hybrids.

Patterns of leaf growth, transpiration and whole-plant water balance in Populus trichocarpa, P. deltoides and their F(1) hybrids were studied during a soil drying cycle. Plant responses were analyzed during three distinct stages of dehydration. In stage I, the transpiration rate of drought-stressed plants remained constant and equal to that of well-watered plants even though soil water content declined by more than 40%. Stage II began as soil and plant water deficits induced stomatal closure. When soil water was expressed as a fraction of transpirable soil water, the transition from stage I to stage II occurred at soil water fractions of 0.35, 0.45 and 0.60 for P. trichocarpa, P. deltoides and their F(1) hybrids, respectively. Reductions in leaf growth coincided with the shift from stage I to stage II. As soil water declined further, decreases in relative transpiration and whole-plant leaf area were significantly greater in parental species than in F(1) hybrids. Inherent feedbacks controlling stomatal water loss and the maintenance and growth of leaf tissue appeared to differ between F(1) and parental genotypes in a pattern characteristic of an overdominant mode of inheritance.Stage III began once the ability of stomata to compensate for water loss had been exhausted. Substantial differences were found in plant survival during stage III, with F(1) hybrids surviving longer than parental species. Survival was more strongly correlated with the hydraulic conductivity of xylem tissues than with the dehydration tolerance of leaf tissues. Collectively, these responses suggest that F(1) hybrids were more drought resistant than either parental species and highlight the importance of whole-plant studies of functional relationships between plant growth, water balance and hydraulic conductivity.

Foliage dark respiration in Abies amabilis (Dougl.) Forbes: variation within the canopy.

Dark respiration of foliage was measured in a 30-year-old stand of Abies amabilis in western Washington from June to November. Both laboratory and field measurements were used to study the effect of environmental and tree variables on respiration. Foliage respiration rates were most strongly influenced by needle temperature. After accounting for leaf temperature differences, foliage respiration decreased with depth in the canopy for all age classes of foliage. Respiration differences attributed to location within the canopy were greatest early in the growing season, but were still significant in November. Younger foliage respired more than older foliage in the upper canopy, but not in the lower canopy. Respiration differences due to foliage age were highly significant in the early growing season, but were not detectable by mid-October.

Crown architecture of Populus clones as determined by branch orientation and branch characteristics.

Crown architecture, including branching pattern, branch characteristics and orientation of proleptic and sylleptic branches was studied in five poplar clones (Populus deltoides, P. trichocarpa and P. trichocarpa x P. deltoides hybrids), grown under intensive culture in the Pacific Northwest, USA. Branch characteristics measured were number, length, diameter, biomass and the angles of origin and termination. The results suggest that genotype has a major influence on crown architecture in Populus. Clonal differences in branch characteristics and branching patterns were found that resulted in striking differences in crown form and architecture. Branch angle and curvature differed significantly among clones, and among height growth increments within clones. Branch length and diameter were significantly correlated in all clones. Sylleptic branches and the considerable leaf area they carry have important implications for whole tree light interception, and thus, play a critical role in the superior growth and productivity of certain hybrid poplar clones. The considerable variation in branch characteristics implies a strong justification for including them in selection and breeding programs for Populus.

Shoot structure, leaf area index and productivity of evergreen conifer stands.

For 12 conifer species, the maximum ratio of shoot to leaf silhouette area of shade-acclimated shoots was found to vary from 0.50 to 0.99. Maximum leaf area index (leaf area per unit ground area) of conifer stands varied from 3.5 to 20, and maximum mean annual increment varied by a factor of 2. Significant correlations were found between leaf silhouette area ratio of shade-acclimated shoots and the maximum leaf area index (R(2) = 0.84) and the maximum mean annual increment (R(2) = 0.93). These results support a hypothesis that species to species differences in the morphology of shade-adapted shoots strongly affect both the development of leaf area and the productivity of stands of evergreen conifers.

Cellular basis for limitation of poplar leaf growth by water deficit.

During the summers of 1986 and 1987, stem and leaf growth were measured on coppiced plants of Populus trichocarpa Torr. & A. Gray, P. deltoides Bartr. ex Marsh, and P. trichocarpa x deltoides growing in the field in Puyallup, WA. The trees were either irrigated periodically throughout the season, or grown without irrigation. In both treatments, stem volume at the end of the growing season was directly proportional to total leaf area in all three genotypes. The rate of individual leaf growth was reduced by lack of irrigation more in the parental species than in the hybrid. Only in the parental species did unirrigated trees have lower leaf water potentials (predawn and midday) than irrigated trees. However, stomatal conductances of all three genotypes were lower in unirrigated trees than in irrigated trees. Osmotic potentials of growing leaves of all three genotypes were also lower in unirrigated trees than in irrigated trees. As a consequence, turgor of growing leaves was as great in unirrigated trees as in irrigated trees, which indicates that turgor differences cannot explain the lower rates of leaf growth in the unirrigated trees. However, cell wall extensibility of leaves was lower in unirrigated trees than in irrigated trees, and the difference was greater in the parental species than in the hybrid. Unlike its effect on leaf area growth, irrigation increased stem volume growth of the hybrid and the parental species by a similar amount (12-16%).

Leaf growth characteristics of fast-growing poplar hybrids Populus trichocarpa x P. deltoides.

Fast-growing hybrid poplar trees (Populus trichocarpa Torr. & A. Gray x P. deltoides Bartr. ex Marsh.) were compared with slower-growing parental types in both field and laboratory experiments to determine physiological components of leaf growth that could be closely related to biomass production. Stem volume was correlated with individual leaf area (r = 0.81) and leaf growth rate (r = 0.82). Hybrids had a greater total leaf area, not because they produced more leaves, but because they had larger leaves than either parental type. The greater leaf size of the hybrids may be explained by inheritance of larger cell number from P. deltoides and larger cell size from P. trichocarpa. Rates of enlargement of isolated leaf discs in liquid culture were approximately 50% of those observed in intact leaves of field-grown plants.

Temperature-Induced Change in the Water Relations of Abies amabilis (Dougl.) Forbes.

Water conductance through Abies amabilis seedlings was measured while the roots were exposed to temperatures from 15 to 0.25 degrees C. Before conductance was measured, the seedlings were preconditioned for 3 months at either a high temperature (23 degrees C) or a low temperature (3 degrees C). For both groups of seedlings, conductance decreased as root temperature decreased. Conductance was lowest at 0.25 degrees C. In addition, preconditioning at 3 degrees C for 3 months significantly lowered conductance to water at all root temperatures. Under the same environmental conditions, seedlings preconditioned at 3 degrees C had less than 25% of the transpirational water loss of seedlings preconditioned at high temperature. A decrease in leaf osmotic potential also resulted from low temperature preconditioning. In trees growing in the subalpine forest, which is the natural habitat of Abies amabilis, both decreased leaf conductance to water vapor and lower osmotic potentials were evident in winter. Since in winter the temperature of the soil in the subalpine zone remains less than 1 degrees C for many months, lowered leaf conductance and decreased osmotic potentials appear to be mechanisms which aid in preventing desiccation damage.

Foliage damage in coniferous trees following volcanic ashfall from Mt. St. Helens.

The foliage of coniferous plants in the area to the northeast of Mt. St. Helens, Washington State, was exposed to heavy ashfall during the May 18, 1980 eruption of the volcano. Significant damage to the pre-1980 foliage occurred after the eruption and continued through the summer. The amount of damage seen on the needles was significantly related to the amount of ash on the foliage.Elevated temperatures caused the foliage damage. The presence of ash on the foliage increased the dimensions of the shoot, thus increasing the boundary layer resistance. In turn this change in geometry elevated needle temperatures. Typical maximum needle temperatures for ash-laden foliage of Abies amabilis were in the range of 35° to 45° C and were 10° C above those of plants without ash. Damage occurred to needles at 40° C after a short growth-chamber exposure. Temperatures within the ash on the foliage also exceeded 40 degrees C.Neither chemical nor mechanical (abrasion) damage occurred. There was no melting of the cuticle. The plants with ash-covered foliage did not experience lower water potentials than those of control plants. The total radiation reflected from the needles was similar for foliage with and without ash.

Drought relations of shrub species: assessment of the mechanisms of drought resistance.

Relatively static factors such as depth of rooting and cuticular conductance and relatively dynamic factors such as stomatol control and changes in the components of water potential were used to assess the drought resistance characteristics of six deciduous shrub species. Predawn water potential during a prolonged drought averaged-2.13 and-3.51 MPa in species known to have deep and shallow patterns of rooting, respectively. It is thus surprising that the osmotic potential at the turgor loss point averaged only-3.01 MPa in the shallow rooted group. The water potential at which irreversible cell damage occurred was the same in both groups (-4.9 MPa), and minimum values observed in the field never dropped below-4.0 MPa. There was, however, a pronounced difference between the two groups with regard to stomatal behavior. This allowed us to characterize the deep-rooted species as avoiders of stress which would cause prolonged stomatal closure whereas the shallow-rooted species had to tolerate prolonged periods of closed stomata.

Net Photosynthesis and Early Growth Trends of a Dominant White Oak (Quercus alba L.).

Examination of the relationship between photosynthesis and growth of a dominant white oak (Quercus alba L.) tree has shown that most growth processes were either completed or well underway before the establishment of significant positive rates of net photosynthesis. Growth was initiated first in the root system (March 3), followed by stem cambial growth (March 26) and later by flower, leaf, and branch growth (April 10). During the period of rapid leaf and branch growth, root and cambial growth ceased and then resumed as the leaves approached maturity. The rapid rate of leaf maturation, the early appearance of positive rates of net photosynthesis in leaves (15% of final size) and the CO(2)-refixing capability of elongating branch tissue reduced the period of time that this white oak tree was dependent on stored reserves. Lower temperature optima and compensation points in developing leaves and stems indicated that the growth-temperature response was optimized for the lower seasonal temperatures observed during the spring. This temperature adaptation further reduced the time that this tree was dependent on stored reserves.

Environmental influences on photosynthesis within the crown of a white oak.

Photosynthesis, leaf conductance, xylem pressure potential as well as six environmental factors were measured within the crown of a dominant, 18.9 m, white oak tree (Quercus alba L.) during the summers of 1974 and 1975. Photosynthesis was measured with a 14CO2 apparatus which enabled a description of the within crown and with leaf variations to be made. Photosynthesis averaged 7.0, 5.6 and 4.6 mgCO2 dm-2 h-1 for top, middle and bottom crown positions. High leaf temperatures and mid-day stomatal closure appeared to limit photosynthesis in the upper crown while light appeared limiting in the lower. Under conditions of a moderate drought, the lower crown's photosynthethic activity approached that of the upper crown. Seasonal shifts in photosynthesis were noted and these were related to shifts in leaf conductance associated with temperature or drought or both pre-conditioning. For example, the xylem pressure potential threshold for stomatal closure varied from-18.9 to-26.1 bars depending upon drought history. With the 14CO2 technique, photosynthetic activity was statistically uniform over the entire leaf surface; thus sampling location within a white oak leaf was not a matter of concern. Similarly, no statistically significant within-leaf variation was found with respect to leaf conductance.