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/ Spatially explicit models that combine remote sensing with geographic information systems (GIS) offer great promise to land managers because they consider the arrangement of landscape elements in time and space. Their visual and geographic nature facilitate the comparison of alternative landscape designs. Among various activities associated with forest management, none cause greater concern than the impacts of timber harvesting on the composition, structure, and function of landscape ecosystems. A timber harvest allocation model (HARVEST) was used to simulate different intensities of timber harvest on 23,592-ha hypothetical landscapes with varying sizes of timber production areas and different initial stand age distributions. Our objectives were to: (1) determine the relative effects of the size of timber production areas, harvest intensity, method used to extract timber, and past timber harvest activity on the production of forest interior and edge; and (2) evaluate how past management (in the form of different initial stand age distributions) constrains future timber production options. Our simulations indicated that the total area of forest interior and the amount of forest edge were primarily influenced by the intensity of timber harvest and the size of openings created by harvest. The size of the largest block of interior forest was influenced most by the size of timber harvests, but the intensity of harvest was also significant, and the size of nontimber production areas was important when harvests were numerous and widely dispersed within timber management areas, as is often the case in managed forests. Stand age-class distributions produced by past harvest activity limited the amount of timber production primarily when group selection was used, but also limited clear-cutting when recent harvest levels were high.KEY WORDS: Simulation modeling; Timber harvest; Historical context; Spatial context; Landscape pattern; Forest interior; Forest edge

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.

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.

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.

Carbon distribution and metabolism by northern red oak seedlings (Quercus rubra L.) were followed for 72 h after a 30-min photosynthetic exposure to (14)CO(2). Approximately 50% of the assimilated carbon was lost during this time, presumably through respiration. Most of the (14)C recovered in the plant remained in the source leaf. Most of the exported (14)C was found in stems and especially roots. Position of the source leaf did not affect distribution of translocated (14)C; however, upper source leaves retained slightly more (14)C than lower source leaves. Most (14)C in all tissues was found initially in sugars. Labeled carbon in this fraction decreased rapidly and increased in other chemical fractions, particularly residue (structural carbohydrates and lignin). More (14)C was incorporated into residue than into any other chemical fraction, indicating continued growth by each of the tissues during the lag stage of seedling development. Labeled carbon increased in proteins for 6 to 12 h after treatment, then remained essentially constant for the remainder of the transport period, indicating both rapid incorporation of the pulsed (14)C into, and slow turnover of, the labeled protein components. In source leaves, (14)C incorporated into starch in the light was lost during the dark period, a typical diurnal storage pattern. In stems, (14)C in starch increased for 12 h, then decreased during the dark period, and then increased during the rest of the transport period. This pattern indicates that stems have both diurnal and long-term storage pools of starch. In roots, (14)C in starch increased rapidly for 12 h, then remained constant for the rest of the transport period, indicating primarily long-term storage in this tissue. Peak (14)C in amino acids in taproot preceded that in stem tissue, indicating recycling of (14)C from the roots to the shoot in amino acids.

Concentrations of glucose, sucrose, soluble reducing sugars, starch and total non-structural carbohydrate were determined during propagation of cuttings from sexually mature Pinus banksiana Lamb. trees. Such cuttings rarely initiate adventitious roots whatever the method or duration of propagation. Terminals, needles, and upper and basal stem segments of cuttings were analyzed at Day 0 and every 2 days for 18 days. Comparison of the results with those of earlier studies with cuttings of P. banksiana seedlings, which root readily, indicated pronounced differences in carbohydrate concentrations and partitioning between the two types of cutting. Compared with those from seedlings, cuttings from sexually mature trees exhibited: (1) more total non-structural carbohydrate in each tissue at Day 0; (2) decreasing rather than increasing total carbohydrate (mainly starch) concentrations in each tissue during propagation; (3) different carbohydrate concentration ratios in each tissue during propagation; and (4) higher sucrose concentrations in terminals during propagation, relative to concentrations at Day 0. Cuttings from sexually mature trees also differed from cuttings of seedlings in having a much lower rate of dry matter accumulation during propagation. These findings suggest that the poor rooting ability of cuttings from sexually mature P. banksiana is not attributable to a lack of total carbohydrate, but that the rooting abilities of cuttings from seedlings and from sexually mature trees differ because of differences between the two types of cutting in rates of net photosynthesis and starch metabolism. The difference in starch metabolism becomes apparent during the first 2 days of propagation.

Diurnal changes in concentrations of leaf chemical fractions and partitioning of photosynthetically fixed (14)C within the plant and among chemical fractions were studied in rapidly growing cottonwood (Populus deltoides Bartr. ex Marsh.) seedlings. During the light period, leaf weight (mg cm(-2)) increased by about 25% primarily as a result of the accumulation of starch and sucrose, and to a lesser extent because of an increase in the content of amino acids and the chloroform fraction (pigments plus lipids). In contrast, reducing sugars and organic acids decreased in concentration. The partitioning of (14)C within the plant also changed during the light period. Acropetal transport to developing leaves and stem decreased from 81 to 55% of the total (14)C translocated from a source leaf in 4 hours, whereas basipetal transport to stem and roots increased from 13 to 37%. Although assimilation rate ((14)C fixed in 0.5 h) remained constant during the light period, the percentage of fixed (14)C translocated out of the source leaf in 4 h decreased from 27 to 9%. This change in transport rate of recently fixed (14)C was caused by a shift in (14)C partitioning from transport sucrose to storage starch. During the light period, the incorporation ratio ((14)C-sugar/(14)C-starch) decreased from 40 at 0700 h to 2 at 1900 h. The partitioning of carbon to different chemical fractions within the source leaf and the interactions or feedback between different sinks and the source leaf have a major influence on plant growth and development. Control of this carbon partitioning is located in both source and sink leaves.