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We studied root growth and respiration of potted plants and field-grown aspen trees (Populus tremuloides Michx.) exposed to ambient or twice-ambient ozone. Root dry weight of potted plants decreased up to 45% after 12 weeks of ozone treatment, and root system respiration decreased by 27%. The ozone-induced decrease in root system respiration of potted plants was more closely correlated with decreased root dry weight than with specific root respiration, suggesting that aspen root metabolism was less affected by ozone than root growth. We used minirhizotrons to study the appearance and disappearance of roots in the field. Length of live roots of field-grown trees increased rapidly early in the season and peaked by midseason in association with a decrease in root production and an increase in root disappearance. In the twice-ambient ozone treatment, live root lengths were 17% less than those of controls, but the effect was not statistically significant. Seasonal soil CO(2) efflux of field-grown trees decreased significantly in the ozone treatments, but because differences in live root length were not significant and root dry weights were not available, the effect on CO(2) efflux could not be attributed directly to decreased root growth.

Clones of aspen (Populus tremuloides Michx.) were identified that differ in biomass production in response to O(3) exposure. (14)Carbon tracer studies were used to determine if the differences in biomass response were linked to shifts in carbon allocation and carbon partitioning patterns. Rooted cuttings from three aspen Clones (216, O(3) tolerant; 271, intermediate; and 259, O(3) sensitive) were exposed to either charcoal-filtered air (CF) or an episodic, two-times-ambient O(3) profile (2x) in open-top chambers. Either recently mature or mature leaves were exposed to a 30-min (14)C pulse and returned to the treatment chambers for a 48-h chase period before harvest. Allocation of (14)C to different plant parts, partitioning of (14)C into various chemical fractions, and the concentration of various chemical fractions in plant tissue were determined. The percent of (14)C retained in recently mature source leaves was not affected by O(3) treatment, but that retained in mature source leaves was greater in O(3)-treated plants than in CF-treated plants. Carbon allocation from source leaves was affected by leaf position, season, clone and O(3) exposure. Recently mature source leaves of CF-treated plants translocated about equal percentages of (14)C acropetally to growing shoots and basipetally to stem and roots early in the season. When shoot growth ceased (August 16), most (14)C from all source leaves was translocated basipetally to stem and roots. At no time did mature source leaves allocate more than 6% of (14)C translocated within the plant to the shoot above. Ozone effects were most apparent late in the season. Ozone decreased the percent (14)C translocated from mature source leaves to roots and increased the percent (14)C translocated to the lower stem. In contrast, allocation from recently mature leaves to roots increased. Partitioning of (14)C among chemical fractions was affected by O(3) more in source leaves than in sink tissue. In source leaves, more (14)C was incorporated into the sugar, organic acid and lipids + pigments fractions, and less (14)C was incorporated into starch and protein fractions in O(3)-treated plants than in CF-treated plants. In addition, there were O(3) treatment interactions between leaf position and clones for (14)C incorporation into different chemical fractions. When photosynthetic data were used to convert percent (14)C transported to the total amount of carbon transported on a mass basis, it was found that carbon transport was controlled more by photosynthesis in the source leaves than proportional changes in allocation to the sinks. Ozone decreased the total amount of carbon translocated to all sink tissue in the O(3)-sensitive Clone 259 because of decreases in photosynthesis in both recently mature and mature source leaves. In contrast, O(3) had no effect on carbon transport from recently mature leaves to lower shoots of either Clone 216 or 271, had no significant effect on transport to roots of Clone 216, and increased transport to roots of Clone 271. The O(3)-induced increase in transport to roots of Clone 271 was the result of a compensatory increase in upper leaf photosynthesis and a relatively greater shift in the percent of carbon allocated to roots. In contrast to those of Clone 271, recently mature leaves of Clone 216 maintained similar photosynthetic rates and allocation patterns in both the CF and O(3) treatments. We conclude that Clone 271 was more tolerant to O(3) exposure than Clone 216 or 259. Tolerance to chronic O(3) exposure was directly related to maintenance of high photosynthetic rates in recently mature leaves and retention of lower leaves.

Rooted cuttings from three aspen (Populus tremuloides Michx.) clones (216, 271 and 259, classified as high, intermediate and low in O(3) tolerance, respectively) were exposed to either diurnal O(3) profiles simulating those of Michigan's Lower Peninsula (episodic treatments), or diurnal square-wave O(3) treatments in open-top chambers in northern Michigan, USA. Ozone was dispensed in chambers ventilated with charcoal-filtered (CF) air. In addition, seedlings were compared to rooted cuttings in their response to episodic O(3) treatments. Early in the season, O(3) caused decreased photosynthetic rates in mature leaves of all clones, whereas only the photosynthetic rates of recently mature leaves of the O(3)-sensitive Clone 259 decreased in response to O(3) exposure. During midseason, O(3) caused decreased photosynthetic rates of both recently mature and mature leaves of the O(3)-sensitive Clone 259, but it had no effect on the photosynthetic rate of recently mature leaves of the O(3)-tolerant Clone 216. Late in the season, however, photosynthetic rates of both recently mature and mature leaves of Clone 216 were lower than those of the control plants maintained in CF air. Ozone decreased the photosynthetic rate of mature leaves of Clone 271, but it increased or had no effect on the photosynthetic rate of recently mature leaves. Photosynthetic response patterns of seedlings to O(3) treatment were similar to those of the clones, but total magnitude of the response was less, perhaps reflecting the diverse genotypes of the seedling population. Early leaf abscission was observed in all clones exposed to O(3); however, Clones 216 and 259 lost more leaf area than Clone 271. By late August, leaf area in the highest O(3) treatment had decreased relative to the controls by 26, 24 and 9% for Clones 216, 259 and 271, respectively. Ozone decreased whole-tree photosynthesis in all clones, and the decrease was consistently less in Clone 271 (23%) than in Clones 216 (56%) and 259 (56%), and was accompanied by declines in total biomass of 19, 28 and 47%, respectively. The relationship between biomass and whole-tree photosynthesis indicates that the negative impact of O(3) on biomass in the clones was determined largely by lower photosynthetic productivity of the foliage, rather than by potential changes in the carbon relations of other plant organs.

We evaluated several sampling schemes for estimating instantaneous whole-tree photosynthesis of 1-year-old Populus clones. Growth of two clones was simulated under varying weather conditions and leaf orientation scenarios providing photosynthetic data on a leaf-by-leaf basis throughout the growing season. Simple random sampling, stratified random sampling and a series of physiologically based sampling schemes were evaluated using either whole-leaf photosynthesis or photosynthetic rate (i.e., photosynthesis per unit leaf area) as the sampling attribute. Ratio and regression estimators with leaf area as an auxiliary variable were also studied. On the basis of their bias and accuracy in estimating instantaneous whole-tree photosynthesis (mg CO(2)), the physiologically based sampling schemes were superior for all combinations of clone type, weather condition and leaf orientation. Aspects of extending the sampling process to estimate daily and seasonal photosynthesis are also elaborated.

Field data from poplar plantations in Michigan, Washington, and Wisconsin were used to validate ECOPHYS, a whole-tree growth process model for juvenile poplar. Five clones representing a range of morphological, phenological, and physiological characteristics were planted on the same date at the three sites. Height and diameter measurements were made monthly on 20 trees per clone, and intensive morphological measurements were made every two weeks on two trees per clone. Hourly solar radiation and temperature data were recorded at each site over the growing season. The model was run for each clone x site combination with the weather data and clonal parameters as inputs. A repeated measures ANOVA indicated that there were significant differences in height growth patterns among both clones and sites, as well as significant clone x site interactions. The model generally predicted height growth within a standard deviation of the field plantation means; three of the 15 clone x site simulations were significantly different from the plantation means. The median error between predicted and observed values was 5%. Evaluation of the clonal parameters showed that differences in photosynthetic rates, morphological attributes such as specific leaf area, and timing of budset are primary factors leading to differences in growth.

Root morphology, biomass, and (14)C distribution were studied in two 2-year-old Populus trichocarpa x P. deltoides hybrids, which originated from hardwood cuttings, to determine the pattern of root distribution in a plantation and to refine methods for root recovery. The trees were labeled with (14)CO(2) and harvested after a 72-hour chase period. Roots attached to each labeled tree were analyzed for morphological traits at the time of harvest. Detached roots from within a 1-m(3) volume of soil surrounding each tree were separated from the soil and sorted on the basis of rooting depth and root diameter. Lateral roots > 2 mm in diameter had a largely horizontal orientation at their point of origin from the cutting and extended horizontally up to 4 m from the cutting. This resulted in considerable overlap of root systems in the plantation. Results from (14)C labeling indicated that 24 +/- 4% (+/- SD) of the carbon exported from branches-labeled within two weeks after branch budset-was translocated to the root system. Dilution of the root (14)C label indicated that from 0 (> 5 mm diameter roots) to 75% (< 2 mm diameter roots) of the roots recovered from within the 1-m(3) volume of soil surrounding a harvested tree originated from other trees. Total root biomass was 6 +/- 1 Mg ha(-1) for both hybrids. Sixty percent of the root biomass was recovered directly from excavation, 16% from coarse-sieving excavated soil, and 24% from re-sorting sieved soil. The study indicated that root growth of hybrid poplars may be rapid and extensive and that detailed sorting of soil subsamples substantially improves the recovery of fine roots < 2 mm in diameter.

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.

A model of photosynthate production is the central component of a larger whole-tree ecophysiological growth process model for Populus (ECOPHYS). This photosynthesis model was validated by comparing predicted photosynthate production values for individual leaves and the total tree with hourly field measurements collected on four days spaced throughout a growing season. Simulated trees had identical numbers of leaves and leaf areas as the sample trees studied in the field, and hourly weather data collected on the plantation site were supplied as a model input. Total production for the four sample days ranged between 200 and 4900 mg CO(2) tree(-1) day(-1). Model predictions of total daily photosynthate production were within 12% of the observed rates for three of the four sampling days. Diurnal variations in stomatal conductance and ambient CO(2) concentrations and seasonal variations in area leaf weight were the primary sources of error. Total leaf area, proportion of sunlit leaf area, and photosynthetic efficiency were the most important factors influencing carbon dioxide exchange rates.

The ECOPHYS model is an ecophysiological growth process model of juvenile poplar clones growing under near optimal conditions. The theoretical basis for the ECOPHYS model is that (1) individual leaves drive and control growth; (2) the microenvironment at the leaf exerts primary control of photosynthetic rates; (3) leaf orientation is a major determinant of that microenvironment, (4) photosynthates produced by leaves are allocated among meristematic and respiratory sinks: and (5) the plant's genome and microenvironment regulate photosynthate allocation. The major driving variables are solar radiation, temperature, and clonal morphological and physiological factors. The user can interact or override any or all of the input variables to examine the effects of such changes on photosynthetic production and growth. Verification and sensitivity analyses of ECOPHYS are presented and discussed. The use of ECOPHYS as a research tool is illustrated with several examples. Model potential and limitations are discussed.

Effects of the contrasting leaf display of poplar cultivars Eugenei (Populus x euramericana) and Tristis (P. tristis x P. balsamifera) on light interception and photosynthesis were studied in the second year of growth in an irrigated plantation near Rhinelander, Wisconsin, USA (lat. 45 degrees N). Leaves on the current terminal (CT) and on proleptic branches were measured between 0900 and 1500 h on five clear days from June to September 1980. Leaf orientation-based differences between these cultivars were evident as the second growing season progressed and the crowns of the trees in the plantation grew together. Leaves of Eugenei are erectophile or tilted from the horizontal. In this cultivar light penetrated throughout the crown; many leaves on the lowest branches were illuminated as fully as those on the upper CT and had higher photosynthetic rates than equivalent leaves in Tristis. However, by early September many of the lower branches on Eugenei trees had abscised. In the planophile Tristis, adaxial photon flux densities (PPFD) of leaves on the lower portion of the CT and on branches were only a fraction of those measured on the upper CT. This pattern became more extreme as the season progressed. Few of the lower branches of Tristis abscised during the growing season. Photosynthesis rates, especially on a whole-leaf basis, were closely related to incident PPFDs in both cultivars. The ecological significance of these results are discussed, as well as the hypothesized effect of leaf inclination on crop productivity.

Diurnal and seasonal photosynthesis patterns were studied in poplar clones Populus tristis Fisch. x P. balsamfera L. cv. Tristis #1 (NC 5260) and Populus x euramericana (Dode) Guiner cv. Eugenei (NC 5326, Carolina poplar) during their first season in the field in a short rotation, intensive culture plantation. Photosynthetic rates were low in immature leaves; increased basipetally on the shoot and peaked in leaves that had recently reached full expansion; and thereafter declined in lower-crown leaves in both clones. Photosynthesis was associated with leaf age and stomatal conductance in immature leaves; adaxial photosynthetic photon flux density (PPFD) and leaf temperature in recently mature leaves; and leaf age and adaxial PPFD in lower-crown leaves. Diurnal photosynthesis patterns within trees were highly variable due to differential light interception among leaves. Results of clonal comparisons of photosynthetic rates were dependent on which leaves were pooled for comparison and how photosynthesis was expressed. Compared to Eugenei, Tristis produced smaller leaves which had higher unit-area photosynthesis rates. The more indeterminate Eugenei outgrew Tristis principally because it more fully utilized the growing season for leaf area production. Photosynthetic production integrated over the growing season was closely related to dry matter production in both clones.

Attempts to relate plant metabolic activity with developmental stage are often hindered by lack of an appropriate developmental index. Existing indices of morphological development are unsuitable for use with plants having a semideterminate, recurrently flushing pattern of growth as displayed by Quercus seedlings. We propose the following morphological index (QMI) to define the stages of Quercus seedling ontogeny: (1) radicle emergence; (2) epicotyl emergence from the soil; and (for each flush) (3) termination of elongation of the second internode, which corresponds with the period of most rapid stem elongation; (4) completion of elongation by all internodes, which corresponds with the period of most rapid leaf elongation; and (5) completion of elongation of the last leaf but one, which usually precedes closely the pause between one growth flush and another. The relationship between QMI and net photosynthesis by individual leaves of Quercus rubra L. seedlings was determined. Net photosynthesis increased with QMI during a flush, but at a particular QMI stage, generally decreased from one flush to the next.

Weekly morphological measurements of trees in permanent growth plots and periodic destructive sampling were used to monitor growth and development of two Populus clones with contrasting morphology and phenology during the establishment year in a short-rotation, intensive-culture system. Tristis (P. tristis Fisch. x P. balsamifera L.) grew rapidly for 48 days before setting bud in July. By contrast, Eugenei (P. x euramericana (Dode) Guinier) grew at a slower rate than Tristis, but maintained this rate for 75 days before setting bud in September. By early October, the total leaf area and dry weight of Eugenei exceeded that of Tristis by 39 and 11%, respectively. In addition, Eugenei had a greater harvest index than Tristis throughout most of the growing season because a larger proportion of photosynthate produced was directed to shoot growth; however, a high shoot/root ratio in Eugenei predisposed it to water stress. Differences in aboveground biomass between clones were largely attributable to clonal differences in seasonal leaf area development.