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Physical mapping and characterization of 5S rRNA genes in Douglas-fir.

Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] is one of the world's most valuable timber species and is widely used in reforestation. Therefore there is considerable interest in understanding its genetic structure. In conifers approximately 75% of the genome is composed of repetitive DNA. Thus for conifers characterization of repetitive DNA is a significant part of genome analysis. We have characterized the organization of 5S rRNA genes in Douglas-fir at both the molecular and chromosome levels. 5S DNA repeat units containing the coding sequence for 5S rRNA and the nontranscribed spacer (NTS) were cloned using PCR. Sequencing and Southern hybridization revealed repeat units of 888 and 871 bp in length, the latter with a 17 bp deletion in the NTS. The coding region showed high homology with other eukaryotic 5S rRNA genes. A 35 bp region of the NTS immediately upstream of the 5' end of the coding region showed high similarity to other conifers but not to other published plant 5S rDNA sequences. Physical mapping of 5S rDNA by fluorescent in situ hybridization using a biotinilated homologous probe revealed a single subtelomeric site on one pair of large metacentric chromosomes.

Base Sequence↗

Ethanol synthesis and aerobic respiration in the laboratory by leader segments of Douglas-fir seedlings from winter and spring.

Stem segments from terminal leaders of Douglas-fir, Pseudotsuga menziesii (Mirb.) Franco, seedlings were sampled in mid-December when cambial cells were dormant. The residual, debudded leaders were resampled again in early May when the cambium was metabolically active. May stems had higher constitutive ethanol concentrations than December stems. This was not the result of cambial hypoxia generated by rapid spring respiration rates, because when aerobic respiration was stimulated by incubating the stems in air at 30 degrees C ethanol production was induced in December, but not in May. Rapid respiration rates at 30 degrees C may have depleted O(2) supplies and induced ethanol production in December stems because dormant, thick-walled cambial cells may be less permeable to CO(2) and O(2), compared with metabolically active, thin-walled cambial cells in May. December stem segments incubated in a N(2) atmosphere at 30 degrees C synthesized 1.8 times more ethanol than segments from May, most likely because spring growth had reduced the soluble sugars available for fermentation. CO(2) efflux from May stems (after 5.5 h of incubation at 30 degrees C) was equal to December stems per unit volume, but greater than December stems per unit surface area. N(2)-induced ethanol concentrations were positively related with CO(2) efflux per unit volume, indicating that rapidly respiring leaders can maintain rapid fermentation rates, provided soluble sugars are readily available. N(2)-induced ethanol and CO(2) efflux per unit volume declined with increasing leader diameter in both seasons, whereas there were no relationships between CO(2) efflux per unit surface area and diameter. Cambium physiology and phenology influence the induction of fermentation and concentrations of ethanol produced in terminal leaders of Douglas-fir, and probably other conifers as well. This needs to be considered when comparing fermentation among species, or comparing individuals from different seasons, or disparate ages within a species.

Aerobiosis↗

Frost injury prediction model for Douglas-fir seedlings in the Pacific Northwest.

Because simple seed- or breeding-zone guidelines are inadequate for controlling the risk of maladaptation to environmental stresses, we are developing operational procedures to assess the risk of frost kill to genetically improved families of Douglas-fir (Pseudotsuga menziesii Mirb. Franco). We have (1) determined the time course of cold hardening and dehardening of nursery-grown Douglas-fir seedlings over four winters, by means of controlled freezing tests, (2) fitted curves to relationships between temperature sum and both fall cold hardening and spring dehardening, (3) applied the temperature sum models to daily temperature records of 80 weather stations to estimate frequency of years with significant frost kill at those stations, (4) interpolated frost kill probabilities for tree farms, using a thin plate spline procedure with elevation, latitude and longitude as variables, and (5) prepared a coarse-scale frost risk map from the resulting grid point estimates. With the exception of a few high-elevation stations, the most damaging frost at any station in any year occurred in either the fall (October and November) or late spring (mid-April to mid-May). In general, damaging spring frosts were two to three times more frequent than fall frosts, and areas in Oregon were at greater risk than areas at similar elevations and longitudes further north. The spline surface was less precise for predicting spring frost risk than fall frost risk.

Journal Article↗

Evidence for red:far red signaling and photomorphogenic growth response in Douglas-fir (Pseudotsuga menziesii) seedlings.

In a greenhouse experiment, potted coastal Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) seedlings were grown in miniature "Nelder" (Nelder 1962) plots where growing space varied from 265 to 2555 cm(2) per plant. After thirty weeks, mean plant height, crown biomass and branch number increased significantly (P = 0.0141) with decreasing growing space (increasing plant density). Differences in height growth became apparent about six weeks after sowing. Furthermore, horizontally reflected radiation measured within the Nelder plots showed a decrease in red:far-red ratio (R:FR) from 1.2 at the lowest density to 0.71 at the highest. Plant height was strongly inversely correlated with estimated phytochrome photoequilibrium values (r(2) = 0.893). Field measurements made in a three-year-old variable density plantation also showed a decrease in R:FR with increasing planting density from 300 to 3,000 trees ha(-1). These results support the hypothesis that young Douglas-fir seedlings are able to detect, through the phytochrome system, the presence of nearby seedlings owing to the depletion of R relative to FR in the spectra reflected by the foliage of the adjacent plants. They then adjust their growth allometry in a way that reduces the possibility of being over topped by these future competitors.

Journal Article↗

Effects of elevated CO(2) and temperature on cold hardiness and spring bud burst and growth in Douglas-fir (Pseudotsuga menziesii).

We examined effects of elevated CO(2) and temperature on cold hardiness and bud burst of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) seedlings. Two-year-old seedlings were grown for 2.5 years in semi-closed, sunlit chambers at either ambient or elevated (ambient + ~ 4 degrees C) air temperature in the presence of an ambient or elevated (ambient + ~ 200 ppm) CO(2) concentration. The elevated temperature treatment delayed needle cold hardening in the autumn and slowed dehardening in the spring. At maximum hardiness, trees in the elevated temperature treatment were less hardy by about 7 degrees C than trees in the ambient temperature treatment. In general, trees exposed to elevated CO(2) were slightly less hardy during hardening and dehardening than trees exposed to ambient CO(2). For trees in the elevated temperature treatments, date to 30% burst of branch terminal buds was advanced by about 6 and 15 days in the presence of elevated CO(2) and ambient CO(2), respectively. After bud burst started, however, the rate of increase in % bud burst was slower in the elevated temperature treatments than in the ambient temperature treatments. Time of bud burst was more synchronous and bud burst was completed within a shorter period in trees at ambient temperature (with and without elevated CO(2)) than in trees at elevated temperature. Exposure to elevated temperature reduced final % bud burst of both leader and branch terminal buds and reduced growth of the leader shoot. We conclude that climatic warming will influence the physiological processes of dormancy and cold hardiness development in Douglas-fir growing in the relatively mild temperate region of western Oregon, reducing bud burst and shoot growth.

Journal Article↗

Conversion of total leaf area to projected leaf area in lodgepole pine and Douglas-fir.

Three definitions of leaf area index (LAI) in the literature have no predictable relationship with each other. Factors were derived for converting total LAI to projected LAI of horizontal leaves and to projected LAI for inclined leaves of lodgepole pine (Pinus contorta Dougl. var. latifolia Engelm.) and coastal Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) to allow comparison of results from different studies. An algorithm was derived to allow determination of these factors based on twig angles and the angles that the foliage subtends with the twig. Allowances were made for both the vertical and horizontal components of projection. The value of the factor for converting total LAI to projected LAI for inclined leaves was 0.229 + 0.0032 for lodgepole pine and 0.230 + 0.0037 for Douglas-fir. Sensitivity analysis established that this conversion factor was more sensitive to differences in vertical angles of the twigs than to twig rotation or foliar arrangement on the twig.

Journal Article↗

Shoot and root vulnerability to xylem cavitation in four populations of Douglas-fir seedlings.

The objectives of this study were to assess the range of genotypic variation in the vulnerability of the shoot and root xylem of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) seedlings to water-stress-induced cavitation, and to assess the trade-off between vulnerability to cavitation and conductivity per unit of stem cross-sectional area (k(s)), both within a species and within an individual tree. Douglas-fir occupies a broad range of environments and exhibits considerable genetic variation for growth, morphology, and drought hardiness. We chose two populations from each of two varieties (the coastal var. menziesii and the interior var. glauca) to represent environmental extremes of the species. Vulnerability curves were constructed for shoots and roots by plotting the percentage loss in conductivity versus water potential. Vulnerability in shoot and root xylem varied genetically with source climate. Stem xylem differed in vulnerability to cavitation between populations; the most mesic population, coastal wet (CW), was the most susceptible of the four populations. In the roots, the most vulnerable population was again CW; the interior wet (IW) population was moderately susceptible compared with the two dry populations, coastal dry (CD) and interior dry (ID). Root xylem was more susceptible to cavitation than stem xylem and had significantly greater k(s). The trade-off between vulnerability to cavitation and k(s), however, was not evident across populations. The most vulnerable population (CW) had a shoot k(s) of 0.534 +/- 0.067 &mgr;mol m(-2) s(-1) MPa(-1), compared with 0.734 +/- 0.067 &mgr;mol m(-2) s(-1) MPa(-1) for the less vulnerable CD stems. In the roots, IW was more vulnerable than ID, but had the same k(s).

Journal Article↗

Effects of nitrogen source on growth and activity of nitrogen-assimilating enzymes in Douglas-fir seedlings.

Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) seedlings were grown for 68 days in a growth chamber in nutrient solutions with ammonium, nitrate or ammonium nitrate as the nitrogen source. Among the nitrogen sources tested, whole-seedling biomass, relative growth rate (RGR), root and shoot elongation, and number of lateral roots, were greatest in seedlings grown with ammonium. In the absence of nitrogen, plant growth and formation of lateral roots were poor. Initially, glutamine synthetase, NAD-glutamate dehydrogenase and aspartate aminotransferase activities were high in young roots and shoots, but all three enzymatic activities decreased after one month of culture. In root apices, glutamine synthetase and aspartate aminotransferase activities were higher than NAD-glutamate dehydrogenase activity. Enzymatic activities were often higher in ammonium-fed seedlings than in seedings supplied with the other forms of nitrogen. Activities of all three enzymes were significantly reduced in seedlings grown in the absence of nitrogen. The beneficial effect of ammonium is discussed on the basis of its involvement in the assimilation pathways of Douglas-fir.

Journal Article↗

Seasonal patterns of photosynthetic light response in Douglas-fir seedlings subjected to elevated atmospheric CO(2) and temperature.

Increases in atmospheric CO(2) concentration and temperature are predicted to increase the light response of photosynthesis by increasing light-saturated photosynthetic rates and apparent quantum yields. We examined the interactive effects of elevated atmospheric CO(2) concentration and temperature on the light response of photosynthesis in Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) seedlings. Seedlings were grown in sunlit chambers controlled to track either ambient (~400 ppm) CO(2) or ambient + 200 ppm CO(2), at ambient temperature or ambient + 4 degrees C. Photosynthetic light response curves were measured over an 18-month period beginning 32 months after treatments were initiated. Light-response curves were measured at the growth CO(2) concentration, and were used to calculate the light-saturated rate of photosynthesis, light compensation point, quantum yield and respiration rate. Elevated CO(2) increased apparent quantum yields during two of five measurement periods, but did not significantly affect light-saturated net photosynthetic rates, light compensation points or respiration rates. Elevated temperature increased all parameters. There were no significant interactions between CO(2) concentration and temperature. We conclude that down-regulation of photosynthesis occurred in the elevated CO(2) treatments such that carbon uptake at a given irradiance was similar across CO(2) treatments. In contrast, increasing temperature may substantially increase carbon uptake rates in Douglas-fir, assuming other environmental factors do not limit photosynthesis; however, it is not clear whether the increased carbon uptake will increase growth rates or be offset by increased carbon efflux through respiration.

Journal Article↗

Coarse root elongation rate estimates for interior Douglas-fir.

Accurate estimates of root growth rates are important for root system modeling, and the spread of root systems may be an important determinant of belowground site occupancy. Estimating root system growth rates is complicated because missing, discontinuous, and false annual growth rings make root cross sections difficult to age. These irregularities can occur even in roots of dominant conifers with rare or absent stem growth ring abnormalities. Incomplete rings were noted in the root growth rings of nine co-dominant interior Douglas-fir (Pseudotsuga menziesii var. glauca (Beissn.) Franco) trees. Coarse root (> 1.0-cm diameter) elongation rates were estimated by fitting a geometric mean regression line to ring count and lateral distance data. In all nine roots examined, the geometric mean regression slope was well within the range of the 95% confidence interval for the ordinary least squares regression of lateral distance versus age, suggesting that measurement error may have been negligible. Coarse root elongation rates (which ranged from 2.8 to 15.3 cm year(-1) and averaged 7.4 cm year(-1)) in the interior Douglas-fir trees studied were much lower than those reported by others. This discrepancy may be a result of limited soil water availability, soil heterogeneity (both soil water content and soil texture were highly variable across short distances) and fragmentation of belowground growing space.

Journal Article↗

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.

Journal Article↗

Importance of needle age and shoot structure on canopy net photosynthesis of balsam fir (Abies balsamea): a spatially inexplicit modeling analysis.

We have developed a spatially inexplicit model of canopy photosynthesis for balsam fir (Abies balsamea (L.) Mill.) that accounts for key processes of light-shoot interaction including irradiance interception by the shoot, spatial aggregation of shoots into branches and crowns, the differential propagation of diffuse and direct light within the canopy, and an ideal representation of penumbra. Also accounted for in the model are the effects of the average radiative climate and shoot age on needle retention, light interception, and photosynthetic capacity. We used reduced versions of this model to quantify the effects of simplifying canopy representation on modeled canopy net photosynthesis. Simplifications explored were the omission of direct beam transformation into penumbral light and the use of different constant shoot properties throughout the canopy. The model was parameterized for a relatively dense balsam fir stand (leaf area index of 5.8) north of Québec City, Canada, and run using hourly meteorological data obtained at the site. The overall performance of the complete model was satisfactory, with maximum values of canopy net photosynthesis of 23 micromol (m(2) ground)(-1) s(-1) (83 mmol m(-2) h(-1)), and a near-saturation of the canopy at a photosynthetically active radiation photon flux density of about 750 micromol m(-2) s(-1) (2.7 mol m(-2) h(-1)). The omission of penumbral effects through the use of unattenuated direct (beam) radiation at all layers of the canopy, as used for broad-leaved species, reduced canopy net photosynthesis by 3.7%. Analysis of the results show that the small impact of penumbra on canopy net photosynthesis stems from the high proportion of diffuse radiation (73%) estimated from our meteorological data set; single-hour results under clear sky conditions approach theoretical bias values of about 30%. Use of mean shoot photosynthetic, light capture and light transmission properties throughout the canopy biased canopy net photosynthesis by less than 3%. However, simulations carried out based on properties of 1-year-old shoots throughout the canopy overestimated canopy net photosynthesis by 9%. Use of the shoot as our smallest functional unit was a potential source of bias because the differential absorption of direct and diffuse radiation within the shoot could not be factored into the model. Other sources of potential bias are discussed.

Abies↗

Mechanisms of Douglas-fir resistance to western spruce budworm defoliation: bud burst phenology, photosynthetic compensation and growth rate.

We compared growth rates among mature interior Douglas-fir (Pseudotsuga menziesii var. glauca (Beissn.) Franco) trees showing resistance or susceptibility to defoliation caused by western spruce budworm (Choristoneura occidentalis Freeman), and among clones and half-sib seedling progeny of these trees in a greenhouse. We also investigated bud burst phenology and photosynthetic responses of clones to budworm defoliation in greenhouse experiments. Resistant mature trees had a higher radial growth rate than susceptible trees, especially during periods of budworm defoliation. Clones from resistant trees grew larger crowns than clones from susceptible trees, whereas stem base diameter at the ground line and height did not differ. Half-sib seedling progeny from resistant trees had larger stem diameter, height, and total biomass than progeny from susceptible trees. Mean 5-year radial growth increment of mature trees was more strongly correlated with growth of seedlings than with growth of clones. Clones from resistant trees had later bud burst than clones from susceptible trees, and budworm defoliation of clones depended on the degree of synchrony between bud burst phenology and budworm larval feeding. Clones of resistant and susceptible mature trees showed similar responses of net photosynthetic rate to 2 years of budworm defoliation. We conclude that phenotypic differences in crown condition of Douglas-fir trees following western spruce budworm defoliation are influenced by tree genotype and that high growth rate and late bud burst phenology promote tree resistance to budworm defoliation.

Animals↗

Effect of feeding by the western conifer seed bug, Leptoglossus occidentalis, on the major storage reserves of developing seeds and on seedling vigor of Douglas-fir.

The impact of feeding by the western conifer seed bug (Leptoglossus occidentalis Heidemann) on storage reserves of developing seeds of Douglas-fir (Pseudostuga menziesii (Mirb.) Franco) was studied by caging nymphal and adult seed bugs on cones during late-season development, and nymphs on cones during early, mid- and late-season development. Analysis of the major storage reserves of partially damaged seeds revealed that late-season feeding by each life stage and feeding by nymphs at all three stages of cone development significantly reduced the amounts of lipid and buffer-insoluble (crystalloid) protein in seeds at harvest by up to 78 and 97%, respectively. Seeds showing light to moderate damage on radiographs did not exhibit a reduction in the amount of buffer-soluble (matrix) protein. Seeds damaged by feeding during early development compensated in part by continuing to synthesize lipid and crystalloid protein. Light or moderate damage to mature Douglas-fir seeds exposed to L. occidentalis in the laboratory reduced seedling emergence by > 80%, but the seedlings that emerged successfully appeared to suffer no adverse effects when grown under standard nursery conditions.

Animals↗

Variation in specific needle area of old-growth Douglas-fir in relation to needle age, within-crown position and epicormic shoot production.

Variation in specific needle area (SNA; cm2 projected fresh needle area g-1 oven-dried needle weight) was investigated in relation to needle age, within-crown position and epicormic shoot production in 450-year-old Douglas-fir (Pseudotsuga menziesii Mirb. (Franco) var. menziesii) trees. Specific needle area decreased with increasing needle age. The magnitude and rate of change in SNA with needle age were greatest for lower-crown branches, and decreased toward the middle- and upper-crown branches. For all branches, there was no difference between regular and epicormic shoots in the relationship between SNA and needle age. Specific needle area decreased with increasing distance from branch base, and this relationship was significant for the majority of needle age classes of the upper- and middle-crown branches. In the lowercrown branches, SNA did not vary with distance from branch base for the majority of needle age classes. For all branches, there was no difference between regular and epicormic shoots in the relationship between SNA and distance from branch base for the majority of needle age classes. These results indicate that renewal of foliage by epicormic shoot production maintains needle quality. Branch SNA increased linearly with decreasing height in the crown at a mean rate of 0.951 +/- 0.110 cm2 g-1 per vertical meter. Total needle area of branches was estimated from total needle dry weight taking into account within-branch variation in SNA. Analyses of allometric relationships between branch size and foliage amount (needle area and needle dry weight) showed that branch length was a better predictor of foliage amount than branch diameter for old Douglas-fir trees. Total needle dry weight and needle area of the sample trees, estimated from branch length and branch height and taking into account vertical within-crown variation in branch SNA, ranged from 42.4 to 154.2 kg and from 246.2 to 816.0 m2 per tree, respectively.

Plant Leaves↗

Biophysical controls of carbon flows in three successional Douglas-fir stands based on eddy-covariance measurements.

We measured net carbon flux (F(CO2)) and net H2O flux (F(H2O)) by the eddy-covariance method at three Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco)-western hemlock (Tsuga heterophylla (Raf.) Sarg.) sites located in the Wind River Valley of southern Washington State, USA. Stands were approximately 20, 40 and 450 years old and measurements were made between June 15 and October 15 of 1998 in the 40- and 450-year-old stands, and of 1999 in the 20- and 450-year-old stands. Our objectives were to determine if there were differences among the stands in: (1) patterns of daytime F(CO2) during summer and early autumn; (2) empirically modeled relationships between local climatic factors (e.g., light, vapor pressure deficit (VPD), soil water content, temperature and net radiation) and daytime F(CO2); and (3) water-use efficiency (WUE). We used the Landsberg equation, a logarithmic power function and linear regression to model relationships between F(CO2) and physical variables. Overall, given the same irradiance, F(CO2) was 1.0-3.9 mol m-2 s-1 higher (P < 0.0001 for both seasons) at the two young stands than at the old-growth stand. During summer and early autumn, F(CO2) averaged 4.2 and 6.1 mol m-2 s-1 at the 20- and 40-year-old stand, respectively. In contrast, the 450-year-old forest averaged 2.2 and 3.2 mol m-2 s-1 in 1998 and 1999, respectively. Increases in VPD were associated with reduced F(CO2) at all three stands, with the greatest apparent constraints occurring at the old-growth stand. Correlations between F(CO2) and all other environmental variables differed among ecosystems, with soil temperature showing a negative correlation and net radiation showing a positive correlation. In the old-growth stand, WUE was significantly greater (P < 0.0001) in the drier summer of 1998 (2.7 mg g-1) than in 1999 (1.0 mg g-1). Although we did not use replicates in our study, the results indicate that there are large differences in F(CO2) among Douglas-fir stands of different ages growing in the same general area, and that variations in age structure and site conditions need to be considered when scaling flux measurements from individual points to the landscape level.

Carbon Dioxide↗

Age- and position-related changes in hydraulic versus mechanical dysfunction of xylem: inferring the design criteria for Douglas-fir wood structure.

We do not know why trees exhibit changes in wood characteristics as a function of cambial age. In part, the answer may lie in the existence of a tradeoff between hydraulic properties and mechanical support. In conifers, longitudinal tracheids represent 92% of the cells comprising the wood and are involved in both water transport and mechanical support. We used three hydraulic parameters to estimate hydraulic safety factors at several vertical and radial locations in the trunk and branches: vulnerability to cavitation; variation in xylem water potential (psi); and xylem relative water content. The hydraulic safety factors for 12 and 88 percent loss of conductivity (S(H12) and S(H88), representing the hydraulic safety factors for the air entry point and full embolism point, respectively) were determined. We also estimated the mechanical safety factor for maximum tree height and for buckling. We estimated the dimensionless hydraulic and mechanical safety factors for six seedlings (4 years old), six saplings (10 years old) and six mature trees (> 110 years old) of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco). Over the natural range of psi, S(H12) decreased linearly from treetop to a minimum of 0.95 at the tree base. Young and mature trees had S(H12) values 1.4 and 1.3 times higher, respectively, at their tips (juvenile wood) than at their bases (mature wood). Modeling analyses indicated that if trees were made entirely of mature wood, S(H12) at the stem base would be only 0.7. The mechanical safety factor was 1.2 times higher for the base of the tree than for the rest of the tree. The minimum mechanical safety factor-1.6 for the critical buckling height and 2.2 for the critical buckling load-occurred at the base of the live crown. Modeling analysis indicated that if trees were made only of mature wood, these values would increase to 1.7 and 2.3, respectively. Hydraulic safety factors had values that were less than half those for mechanical safety factors, suggesting that wood structure in Douglas-fir has evolved primarily as a result of selection for hydraulic safety rather than mechanical safety. The results suggest that forest managers must consider the role of juvenile wood in tree physiology to avoid producing plantations vulnerable to drought.

Oregon↗

Growth response of Douglas-fir seedlings to nitrogen fertilization: importance of Rubisco activation state and respiration rates.

High foliar nitrogen concentration ([N]) is associated with high rates of photosynthesis and thus high tree productivity; however, at excessive [N], tree productivity is reduced. Reports of excessive [N] in the Douglas-fir forests of the Oregon Coast Range prompted this investigation of growth and needle physiological responses to increasing foliar N concentrations in 1-year-old Douglas-fir seedlings. After 1 year of N fertilization, total seedling biomass increased with each successive increase in N fertilizer concentration, except in the highest N fertilization treatment. Of the many physiological responses that were analyzed, only photosynthetic capacity (i.e., Vcmax), respiration rates and leaf specific conductance (KL) differed significantly between N treatments. Photosynthetic capacity showed a curvilinear relationship with foliar [N], reaching an apparent maximum rate when needle N concentrations exceeded about 12 mg g(-1). In vitro measurements of ribulose-1,5-bisphosphate carboxylase (Rubisco) activity suggested that photosynthetic capacity was best related to activated, not total, Rubisco content. Rubisco activation state declined as foliar [N] increased, and based on its significant correlation (r2= 0.63) with foliar Mn:Mg ratios, it may be related to Mn inactivation of Rubisco. Respiration rates increased linearly as foliar N concentration increased (r2= 0.84). The value of K(L) also increased as foliar [N] increased, reaching a maximum when foliar [N] exceeded about 10 mg g(-1). Changes in K(L) were unrelated to changes in leaf area or sapwood area because leaf area to sapwood area ratios remained constant. Cumulative effects of the observed physiological responses to N fertilization were analyzed by modeling annual net CO2 assimilation (Anet) based on treatment specific values of Vcmax, dark respiration (Rdark) and KL. Estimates of Anet were highly correlated with measured total seedling biomass (r2= 0.992), suggesting that long-term, cumulative effects of maximum Rubisco carboxylation, Rdark and KL responses to N fertilization may limit seedling production when foliar N exceeds about 13 mg g(-1) or is reduced to less than about 11 mg g(-1).

Cell Respiration↗