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Trichloroethylene (TCE) is a widespread environmental pollutant common in groundwater plumes associated with industrial manufacturing areas. We had previously isolated and characterized a natural bacterial endophyte, Enterobacter sp. strain PDN3, of poplar trees, that rapidly metabolizes TCE, releasing chloride ion. We now report findings from a successful three-year field trial of endophyte-assisted phytoremediation on the Middlefield-Ellis-Whisman Superfund Study Area TCE plume in the Silicon Valley of California. The inoculated poplar trees exhibited increased growth and reduced TCE phytotoxic effects with a 32% increase in trunk diameter compared to mock-inoculated control poplar trees. The inoculated trees excreted 50% more chloride ion into the rhizosphere, indicative of increased TCE metabolism in planta. Data from tree core analysis of the tree tissues provided further supporting evidence of the enhanced rate of degradation of the chlorinated solvents in the inoculated trees. Test well groundwater analyses demonstrated a marked decrease in concentration of TCE and its derivatives from the tree-associated groundwater plume. The concentration of TCE decreased from 300 μg/L upstream of the planted area to less than 5 μg/L downstream of the planted area. TCE derivatives were similarly removed with cis-1,2-dichloroethene decreasing from 160 μg/L to less than 5 μg/L and trans-1,2-dichloroethene decreasing from 3.1 μg/L to less than 0.5 μg/L downstream of the planted trees. 1,1-dichloroethene and vinyl chloride both decreased from 6.8 and 0.77 μg/L, respectively, to below the reporting limit of 0.5 μg/L providing strong evidence of the ability of the endophytic inoculated trees to effectively remove TCE from affected groundwater. The combination of native pollutant-degrading endophytic bacteria and fast-growing poplar tree systems offers a readily deployable, cost-effective approach for the degradation of TCE, and may help mitigate potential transfer up the food chain, volatilization to the atmosphere, as well as direct phytotoxic impacts to plants used in this type of phytoremediation.

Three young northern temperate forest communities in the north-central United States were exposed to factorial combinations of elevated carbon dioxide (CO2 ) and tropospheric ozone (O3 ) for 11 years. Here, we report results from an extensive sampling of plant biomass and soil conducted at the conclusion of the experiment that enabled us to estimate ecosystem carbon (C) content and cumulative net primary productivity (NPP). Elevated CO2 enhanced ecosystem C content by 11%, whereas elevated O3 decreased ecosystem C content by 9%. There was little variation in treatment effects on C content across communities and no meaningful interactions between CO2 and O3 . Treatment effects on ecosystem C content resulted primarily from changes in the near-surface mineral soil and tree C, particularly differences in woody tissues. Excluding the mineral soil, cumulative NPP was a strong predictor of ecosystem C content (r(2) = 0.96). Elevated CO2 enhanced cumulative NPP by 39%, a consequence of a 28% increase in canopy nitrogen (N) content (g N m(-2) ) and a 28% increase in N productivity (NPP/canopy N). In contrast, elevated O3 lowered NPP by 10% because of a 21% decrease in canopy N, but did not impact N productivity. Consequently, as the marginal impact of canopy N on NPP (∆NPP/∆N) decreased through time with further canopy development, the O3 effect on NPP dissipated. Within the mineral soil, there was less C in the top 0.1 m of soil under elevated O3 and less soil C from 0.1 to 0.2 m in depth under elevated CO2 . Overall, these results suggest that elevated CO2 may create a sustained increase in NPP, whereas the long-term effect of elevated O3 on NPP will be smaller than expected. However, changes in soil C are not well-understood and limit our ability to predict changes in ecosystem C content., (© 2014 The Authors Global Change Biology Published by John Wiley & Sons Ltd.)

Human activity causes increasing background concentrations of the greenhouse gases CO2 and O3. Increased levels of CO2 can be found in all terrestrial ecosystems. Damaging O3 concentrations currently occur over 29% of the world's temperate and subpolar forests but are predicted to affect fully 60% by 2100 (ref. 3). Although individual effects of CO2 and O3 on vegetation have been widely investigated, very little is known about their interaction, and long-term studies on mature trees and higher trophic levels are extremely rare. Here we present evidence from the most widely distributed North American tree species, Populus tremuloides, showing that CO2 and O3, singly and in combination, affected productivity, physical and chemical leaf defences and, because of changes in plant quality, insect and disease populations. Our data show that feedbacks to plant growth from changes induced by CO2 and O3 in plant quality and pest performance are likely. Assessments of global change effects on forest ecosystems must therefore consider the interacting effects of CO2 and O3 on plant performance, as well as the implications of increased pest activity.

Predicting ozone-induced reduction of carbon sequestration of forests under elevated tropospheric ozone concentrations requires robust mechanistic leaf-level models, scaled up to whole tree and stand level. As ozone effects depend on genotype, the ability to predict these effects on forest carbon cycling via competitive response between genotypes will also be required. This study tests a process-based model that predicts the relative effects of ozone on the photosynthetic rate and growth of an ozone-sensitive aspen clone, as a first step in simulating the competitive response of genotypes to atmospheric and climate change. The resulting composite model simulated the relative above ground growth response of ozone-sensitive aspen clone 259 exposed to square wave variation in ozone concentration. This included a greater effect on stem diameter than on stem height, earlier leaf abscission, and reduced stem and leaf dry matter production at the end of the growing season. Further development of the model to reduce predictive uncertainty is discussed.

Atmospheric chemical composition affects foliar chemical composition, which in turn influences the dynamics of both herbivory and decomposition in ecosystems. We assessed the independent and interactive effects of CO2 and O3 fumigation on foliar chemistry of quaking aspen (Populus tremuloides) and paper birch (Betula papyrifera) at a Free-Air CO2 Enrichment (FACE) facility in northern Wisconsin. Leaf samples were collected at five time periods during a single growing season, and analyzed for nitrogen. starch and condensed tannin concentrations, nitrogen resorption efficiencies (NREs), and C:N ratios. Enriched CO2 reduced foliar nitrogen concentrations in aspen and birch; O3 only marginally reduced nitrogen concentrations. NREs were unaffected by pollution treatment in aspen, declined with 03 exposure in birch, and this decline was ameliorated by enriched CO2. C:N ratios of abscised leaves increased in response to enriched CO2 in both tree species. O3 did not significantly alter C:N ratios in aspen, although values tended to be higher in + CO2 + O3 leaves. For birch, O3 decreased C:N ratios under ambient CO2 and increased C:N ratios under elevated CO2. Thus, under the combined pollutants, the C:N ratios of both aspen and birch leaves were elevated above the averaged responses to the individual and independent trace gas treatments. Starch concentrations were largely unresponsive to CO2 and O3 treatments in aspen. but increased in response to elevated CO2 in birch. Levels of condensed tannins were negligibly affected by CO2 and O3 treatments in aspen, but increased in response to enriched CO2 in birch. Results from this work suggest that changes in foliar chemical composition elicited by enriched CO2 are likely to impact herbivory and decomposition, whereas the effects of O3 are likely to be minor, except in cases where they influence plant response to CO2.

To determine whether elevated CO2 reduces or exacerbates the detrimental effects of O3 on aspen (Populus tremuloides Michx.). aspen clones 216 and 271 (O3 tolerant), and 259 (O3 sensitive) were exposed to ambient levels of CO2 and O3 or elevated levels of CO2, O3, or CO2 + O3 in the FACTS II (Aspen FACE) experiment, and physiological and molecular responses were measured and compared. Clone 259. the most O3-sensitive clone, showed the greatest amount of visible foliar symptoms as well as significant decreases in chlorophyll, carotenoid, starch, and ribulose-1, 5-bisphosphate carboxylase/oxygenase (Rubisco) concentrations and transcription levels for the Rubisco small subunit. Generally, the constitutive (basic) transcript levels for phenylalanine ammonialyase (PAL) and chalcone synthase (CHS) and the average antioxidant activities were lower for the ozone sensitive clone 259 as compared to the more tolerant 216 and 271 clones. A significant decrease in chlorophyll a, b and total (a + b) concentrations in CO2, O3, and CO2 + O3 plants was observed for all clones. Carotenoid concentrations were also significantly lower in all clones; however. CHS transcript levels were not significantly affected, suggesting a possible degradation of carotenoid pigments in O3-stressed plants. Antioxidant activities and PAL and 1-aminocyclopropane-l-carboxylic acid (ACC)-oxidase transcript levels showed a general increase in all O3 treated clones, while remaining low in CO2 and CO2 + O3 plants (although not all differences were significant). Our results suggest that the ascorbate-glutathione and phenylpropanoid pathways were activated under ozone stress and suppressed during exposure to elevated CO2. Although CO2 + O2 treatment resulted in a slight reduction of O3-induced leaf injury, it did not appear to ameliorate all of the harmful affects of O3 and, in fact. may have contributed to an increase in chloroplast damage in all three aspen clones.

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.

The ability of a developing cottonwood (Populus deltoides Bartr.) leaf to export (14)C-labeled assimilates begins at the lamina tip and progresses basipetally with increasing LPI. This progression indicates that portions of leaves function quasi-independently in their ability to export (14)C-photosynthate. Although most of the exported radioactivity was recovered in the petiole as water-80% alcohol-soluble compounds, there was also substantial incorporation into the chloroform and insoluble fractions. This observation indicates that assimilates translocated from the lamina are used in structural development of the petiole. Freeze substitution and epoxy embedding were used to prepare microautoradiographs for localization of water-soluble compounds. Radioactivity was found in all cell types within specific subsidiary bundles of the petiole. However, radioactive assimilates appeared to move from the translocation pathway in the phloem toward active sinks in the walls of the expanding metaxylem cells. Translocation in the mature xylem vessels was not observed.

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.

Individual leaves of eastern cottonwood (Populus deltoides), representing an ontogenetic series from leaf plastochron index 0.0 to 8.0, were fed (14)CO2 photosynthetically and then harvested at times ranging from 15 to 1440 min. The lamina of each fed leaf was sectioned from tip to base into 5 parts, and each part was quantitatively assayed for (14)C activity. In young leaves, the percentage of the total (14)C fixed (expressed in dpm/mg of dry leaf tissue) was high in the lamina tip and decreased almost linearly toward the base. With increasing leaf age, the percentage of (14)C fixed decreased in the lamina tip and increased in the base. The relative activity in mature leaves was almost uniform throughout the lamina. No differences were detected in the (14)C distribution patterns within leaves over the time series.On the basis of the data presented and of anatomical observations of developing cottonwood leaves, the hypothesis that the precociously mature lamina tip may provide photosynthates to the still-expanding lamina base was shown to be invalid. It is concluded that bidirectional transport in a developing cottonwood leaf results from simultaneous import to the immature basal region and export from the mature tip.