Carbon dioxide enrichment improves growth, water relations and survival of droughted honey mesquite (Prosopis glandulosa) seedlings

Low water availability reduces the establishment of the invasive shrub Prosopis on some grasslands. Water deficit survival and traits that may contribute to the postponement or tolerance of plant dehydration were measured on seedlings of P. glandulosa Torr. var. glandulosa (honey mesquite) grown at CO(2) concentrations of 370 (ambient), 710, and 1050 micro mol mol(-1). Because elevated CO(2) decreases stomatal conductance, the number of seedlings per container in the elevated CO(2) treatments was increased to ensure that soil water content was depleted at similar rates in all treatments. Seedlings grown at elevated CO(2) had a greater root biomass and a higher ratio of lateral root to total root biomass than those grown at ambient CO(2) concentration; however, these seedlings also shed more leaves and retained smaller leaves. These changes, together with a reduced transpiration/leaf area ratio at elevated CO(2), may have contributed to a slight increase in xylem pressure potentials of seedlings in the 1050 micro mol mol(-1) CO(2) treatment during the first 37 days of growth (0.26 to 0.40 MPa). Osmotic potential was not affected by CO(2) treatment. Increasing the CO(2) concentration to 710 and 1050 micro mol mol(-1) more than doubled the percentage survival of seedlings from which water was withheld for 65 days. Carbon dioxide enrichment significantly increased survival from 0% to about 40% among seedlings that experienced the lowest soil water content. By increasing seedling survival of drought, rising atmospheric CO(2) concentration may increase abundance of P. glandulosa on grasslands where low water availability limits its establishment.     

Physiological Characteristics and Carbohydrate Contents of Spring-lifted Picea glauca Bareroot Seedlings Following Low-temperature Storage

This study examined the effects of low-temperature storage of white spruce [ Picea glauca (Moench) Voss] bareroot seedlings to determine whether the time between lifting and planting of spring-lifted seedlings could be extended. Seedlings were lifted from the nursery beds on May 4, 1994, and stored at- 2C (frozen storage) and 4C (cold storage) for 3, 5 and 7 weeks. Frozen storage of spring-lifted seedlings resulted in an increase in sugar levels in roots and needles that progressed with storage duration. Seedlings stored in cold storage maintained a relatively constant sugar content. Needle starch content decreased with storage duration in both frozen and cold storage. In the roots of cold-stored seedlings, starch content remained relatively constant during storage; however, in the frozen-stored seedlings, root starch levels sharply declined during the initial 3 weeks of storage. The levels of total non-structural carbohydrates (starch and sugars) decreased in both types of storage in needles, but not in roots. However, the decrease was more pronounced in the cold-stored than in the frozen-stored seedlings. Gas exchange, root growth potential and number of days to bud break were similar in frozen- and cold-stored seedlings planted in the greenhouse. However, following planting in the forest, cold-stored seedlings flushed buds earlier than did frozen-stored seedlings. The results indicate that tree nurseries could consider frozen storage of spring-lifted white spruce seedlings to facilitate lifting and planting schedules.     

Effects of soil temperature on biomass and carbohydrate allocation in Scots pine (Pinus sylvestris) seedlings at the beginning of the growing season

We studied effects of soil temperature on shoot and root extension growth and biomass and carbohydrate allocation in Scots pine (Pinus sylvestris L.) seedlings at the beginning of the growing season. One-year-old Scots pine seedlings were grown for 9 weeks at soil temperatures of 5, 9, 13 and 17 degrees C and an air temperature of 17 degrees C. Date of bud burst, and the elongation of shoots and roots were monitored. Biomass of current and previous season roots, stem and needles was determined at 3-week intervals. Starch, sucrose, glucose, fructose, sorbitol and inositol concentrations were determined in all plant parts except new roots. The timing of both bud burst and the onset of root elongation were unaffected by soil temperature. At Week 9, height growth was reduced and root extension growth was much less at a soil temperature of 5 degrees C than at higher soil temperatures. Total seedling biomass was lowest in the 5 degrees C soil temperature treatment and highest in the 13 degrees C treatment, but there was no statistically significant difference in total biomass between seedlings grown at 13 and 17 degrees C. In response to increasing soil temperature, below-ground biomass increased markedly, resulting in a slightly higher allocation of biomass to below-ground parts. Among treatments, root length was greatest at a soil temperature of 17 degrees C. The sugar content of old roots was unaffected by soil temperature, but the sugar content of new needles increased with increasing soil temperature. The starch content of all seedling parts was lowest in seedlings grown at 17 degrees C. Otherwise, soil temperature had no effect on seedling starch content.     

Growth rate and survivorship of drought: CO2 effects on the presumed tradeoff in seedlings of five woody legumes

Traits that promote rapid growth and seedling recruitment when water is plentiful may become a liability when seedlings encounter drought. We tested the hypothesis that CO2 enrichment reinforces any tradeoff between growth rate and drought tolerance by exaggerating interspecific differences in maximum relative growth rate (RGR) and survivorship of drought among seedlings of five woody legumes. We studied invasive species of grasslands that differ in distribution along a rainfall gradient. Survivorship of drought at ambient CO2 concentration ([CO2]) was negatively related to RGR in well-watered seedlings in one of two experiments, but the relationship was weak because interspecific differences in RGR were small. Contrary to our hypothesis, there was no significant relationship among well-watered seedlings between RGR at ambient [CO2] and either the relative or absolute increase in RGR at elevated [CO2]. As predicted, however, CO2 enrichment reinforced interspecific differences in survivorship of seedlings exposed to similar rates of soil water depletion. Doubling [CO2] improved seedling survivorship of the most drought-tolerant species throughout the period of soil water depletion, but did not consistently affect survivorship of more drought-sensitive species. Midday xylem pressure potentials of drought-treated seedlings were less negative at elevated [CO2] than at ambient [CO2], but no other measured trait was consistently correlated with improved survivorship at high [CO2]. Carbon dioxide enrichment may not reinforce species differences in RGR, but could exaggerate interspecific differences in drought tolerance. To the extent that seedling persistence in grasslands correlates with drought survivorship, our results indicate a positive effect of CO2 enrichment on recruitment of woody legumes that are currently tolerant of drought.     

Responses of loblolly pine seedlings to elevated CO(2) and fluctuating water supply

Osmotic adjustment of loblolly pine (Pinus taeda L.) seedlings to fluctuating water supply in elevated CO(2) was investigated. Seedlings were grown in controlled-environment chambers in either 350 or 700 micro l l(-1) CO(2) with weekly watering for four months, after which they were either watered weekly (well-watered treatment) or every two weeks (water-stress treatment) for 59 days. Osmotic adjustment was assessed by pressure-volume analysis of shoots and by analysis of soluble carbohydrates and free amino acids in roots during the last drying cycle. In well-watered seedlings, elevated CO(2) increased the concentration of soluble sugars in roots by 68%. Water stress reduced the soluble sugar concentration in roots of seedling growing in ambient CO(2) to 26% of that in roots of well-watered seedlings. Elevated CO(2) mitigated the water stress-induced decrease in the concentration of soluble sugars in roots. However, this was probably due, in part, to carbohydrate loading during the first four months when all seedlings were grown in the presence of a high water supply, rather than to osmotic adjustment to water stress. Water stress caused a doubling in the concentration of free primary amino acids in roots, whereas elevated CO(2) reduced primary amino acid and nitrogen concentrations to 32 and 74%, respectively, of those in roots of seedlings grown in ambient CO(2). There was no indication of large-scale osmotic adjustment to water stress or that elevated CO(2) enhanced osmotic adjustment in loblolly pine.     

Effect of elevated CO2 on monoterpene emission of young Quercus ilex trees and its relation to structural and ecophysiological parameters

We investigated growth, leaf monoterpene emission, gas exchange, leaf structure and leaf chemical composition of 1-year-old Quercus ilex L. seedlings grown in ambient (350 microl l(-1)) and elevated (700 microl l(-1)) CO2 concentrations ([CO2]). Monoterpene emission and gas exchange were determined at constant temperature and irradiance (25 degrees C and 1000 micromol m(-2) s(-1) of photosynthetically active radiation) at an assay [CO2] of 350 or 700 microl l(-1). Measurements were made on intact shoots after the end of the growing season between mid-October and mid-February. On average, plants grown in elevated [CO2] had significantly increased foliage biomass (about 50%). Leaves in the elevated [CO2] treatment were significantly thicker and had significantly higher concentrations of cellulose and lignin and significantly lower concentrations of nitrogen and minerals than leaves in the ambient [CO2] treatment. Leaf dry matter density and leaf concentrations of starch, soluble sugars, lipids and hemi-cellulose were not significantly affected by growth in elevated [CO2]. Monoterpene emissions of seedlings were significantly increased by elevated [CO2] but were insensitive to short-term changes in assay [CO2]. On average, plants grown in elevated [CO2] had 1.8-fold higher monoterpene emissions irrespective of the assay [CO2]. Conversely, assay [CO2] rapidly affected photosynthetic rate, but there was no apparent long-term acclimation of photosynthesis to growth in elevated [CO2]. Regardless of growth [CO2], photosynthetic rates of all plants almost doubled when the assay [CO2] was switched from 350 to 700 microl l(-1). At the same assay [CO2], mean photosynthetic rates of seedlings in the two growth CO2 treatments were similar. The percentage of assimilated carbon lost as monoterpenes was not significantly altered by CO2 enrichment. Leaf emission rates were correlated with leaf thickness, leaf concentrations of cellulose, lignin and nitrogen, and total plant leaf area. In all plants, monoterpene emissions strongly declined during the winter independently of CO2 treatment. The results are discussed in the context of the acquisition and allocation of resources by Q. ilex seedlings and evaluated in terms of emission predictions.     

Shoot and root sensitivity of containerized black spruce,white spruce and jack pine seedlings to late fall freezing

Damage to containerized forest seedlings due to freezing can occur in the fall or early winter in Canadian forest nurseries. The following spring, damage to shoots and impairment of growth is observed. The objectives of this experiment were to measure the impact of late fall low temperatures (0° to --30°C) on whole seedlings of the three most common species used for reforestation in Quebec: black spruce (BS), white spruce (WS) and jack pine (JP). Impacts of freezing temperatures on (i) whole seedling and apical bud mortality, (ii) shoot growth and root mortality, (iii) stem electrical resistance, (iv) shoot and root water relations, (v) concentrations of N, P, K, Ca, Mg, and total sugars in shoots were assessed. JP showed the highest rate of whole seedling mortality while WS showed the highest rate of apical bud mortality. JP was the most severely affected: destruction of the root system at low temperatures as well as a reduction of shoot growth and stem diameter and a decrease (more negative) in shoot and root water potential. WS showed a reduction of shoot growth despite no apparent damage to the root system at low temperatures. BS was not affected by temperatures as low as --30°C. Nutrient and sugar concentrations were not affected by low temperature treatments.     

Carbon and phosphorus partitioning in Pinus serotina seedlings growing under hypoxic and low-phosphorus conditions

Ten-week-old pond pine (Pinus serotina Michx.) seedlings were grown in solution culture at 5 or 100 microM P and under aerobic or hypoxic solution conditions. After 6 and 10 weeks in the treatments, changes in relative growth rate (RGR), P acquisition and allocation, and carbohydrate partitioning were determined by analyzing tissue for total P, soluble sugars and starch. Six weeks of low-P growth conditions decreased seedling dry weight and the ratio of shoot dry weight to root dry weight (S/R) by 39 and 51%, respectively, in comparison to seedlings from the aerobic, high-P (control) treatment. Mean RGRs of shoots in the low-P treatment were reduced by 33%, whereas root growth was unaffected. After 10 weeks of low-P growth conditions, however, both shoot and root RGRs were significantly reduced, and plants had lower S/R ratios than in any other treatment. Slowed shoot growth was accompanied by starch and nonstructural carbohydrate accumulation in needles, indicating that needle growth was not limited by carbohydrate supply. Six weeks of low-P growth conditions decreased total seedling P by 75%, reflecting a 97% reduction in the net uptake rate (NUR). Shoot NUR as a fraction of seedling NUR was also greatly reduced in the low-P treatment, indicating that low-P growth conditions affected P translocation to the shoot more than P accumulation by roots. In contrast, 6 weeks of hypoxic growth conditions decreased total dry weight of seedlings in the high-P treatment by 41% relative to their aerobic counterparts. Root growth was affected more than shoot growth, however, and S/R ratios increased. After 10 weeks, S/R ratios doubled, primarily because of the reduction in root RGR. Nevertheless, roots of hypoxic seedlings contained a higher percentage of total seedling P than their aerobic counterparts. Net P acquisition per seedling decreased by more than 50% under hypoxic growth conditions, as a result of reductions in both root RGR and seedling NUR. Starch accumulation in shoots of hypoxic seedlings reflected reductions both in root growth and in transport of carbohydrates to nonwoody roots. Carbohydrate availability did not appear to be limiting growth of hypoxic woody roots, which are well-aerated internally, but it may have limited metabolic processes in nonwoody roots of seedlings from the high-P treatment.     

Long- and short-term flooding effects on survival and sink-source relationships of swamp-adapted tree species

About 95% of swamp tupelo (Nyssa sylvatica var. biflora (Walt.) Sarg.) and sweetgum (Liquidambar styraciflua L.) seedlings survived continuous root flooding for more than two years, whereas none of the swamp chestnut oak (Quercus michauxii Nutt.) and cherrybark oak (Q. falcata var. pagodifolia Ell.) seedlings survived one year of flooding. Death of oak seedlings occurred in phases associated with periods of major vegetative growth, e.g., after bud burst in spring, after summer stem elongation, and during the winter deciduous stage, suggesting that stored reserves and sources were inadequate to maintain the seedlings when vegetative sinks were forming. Additional evidence that flooding induced a source deficiency in oak was that leaves of flooded oak were 65 to 75% smaller than leaves of nonflooded oak. Flooded swamp tupelo seedlings had a normal leaf size and patchy stomatal opening compared with nonflooded seedlings. Flooding caused increases in alcohol dehydrogenase (ADH) specific activity in taproot cambial tissues and increases in starch concentrations of swamp tupelo seedlings that were reversed when seedlings were removed from flooding. Flooding had little effect on soluble sugar concentrations in swamp tupelo or sweetgum. In the long-term flood-dry-flood treatment, in which all species had survivors, upper canopy leaf photosynthetic rates were higher in all species during the dry period than in nonflooded controls, whereas their starch and soluble sugars concentrations were similar to those of nonflooded controls. Based on seedling survival and the sink-source relationships, the order of flood tolerance was: swamp tupelo > sweetgum > swamp chestnut oak > cherrybark oak.     

First-year growth response of cold-stored,nursery-grown aspen planting stock

This research examined the first year growth characteristics of cold stored and transplanted nursery-produced aspen (Populus tremuloides) seedlings (container and bareroot (BR)) and compared it to the growth of seedlings that had not been transplanted (established from germinants in the field) and therefore had an unrestricted root system (UR). Prior to planting, nursery-produced seedlings were placed in cold storage (−3°C) and root growth potential (RGP) and total non-structural carbohydrate (TNC) root reserves were tested at 0, 10, 75 and after 150 (container) and 190 days (BR) of storage. Both container and BR stock had much lower root to shoot ratios (RSRs) and root carbohydrate reserves compared to UR seedlings after 170 days. During storage, root reserves in container stock declined faster than in the BR and UR seedlings. RGP in all nursery stock was the highest after 75 days of storage, while longer storage resulted in shoot dieback and reduced root growth. After the first growing season, UR seedlings were one tenth the size of the nursery stock; however, in the second growing season they had no stem dieback and grew twice the height and stem diameter. The higher RSRs and root reserves in the UR seedlings was likely caused by early bud set in its first year of growth. This suggests that inducing bud set earlier in the growing regime might allow seedlings to increase root mass and carbohydrate reserves.     

Influence of nutrient supply on spring frost hardiness and time of bud break in Norway spruce (Picea abies (L.) Karst.) seedlings

When spring frosts occur on recently planted forest sites, severe damage may occur to the seedlings. The aim of the present study was to test how different low levels of nutrient concentrations in Norway spruce (Picea abies (L.) Karst.) seedlings affected spring frost hardiness and time of bud break. Seedlings were grown in a greenhouse for one season and supplied with fertiliser containing 22, 43 and 72 mg N l^–1, respectively. The treatments resulted in needle nitrogen concentrations ranging from 0.9 to 1.8% in autumn. After winter storage at 0 °C, bud break was recorded on seedlings growing in the greenhouse, outdoors and in growth chambers at 12 °C and at 17 °C. Freezing tests were performed on seedlings directly removed from winter storage and following one week growth in the greenhouse. Seedlings receiving fertiliser with 43 mg N l^–1 had less freezing injury than the two other fertilisation treatments in the present study. The earliest bud break occurred in seedlings receiving 72 mg N l^–1.     

Slash pine bud dormancy as affected by lifting date and root wrenching in the nursery

Bud dormancy of root wrenched and unwrenched slash pine (Pinus elliottii Engelm.) seedlings growing in a forest nursery was measured on five lifting dates. Determination of bud dormancy was based on days to budbreak (DBB) under optimal growing conditions, mitotic activity in the apical meristem, chilling hours accumulated, and bud morphology. Based on DBB, seedlings were most dormant at Lift 2 on November 24 after exposure to 189 hours below 10 degrees C and 93 hours below 6.7 degrees C. Mitotic activity in the apical meristem was at its lowest 23 days later at Lift 3, possibly indicating the period when seedlings are most resistant to transplanting stresses. Multiple wrenching resulted in a slight shift in the dormancy cycle as wrenched seedlings set bud sooner in the nursery and broke bud sooner at the planting site in the spring than control seedlings. This implies that wrenched seedlings can be successfully lifted from the nursery earlier and will initiate spring shoot growth earlier than control seedlings.     

Germination and seedling growth in Abies lasiocarpa (Hook.) Nutt. as affected by provenance,seed pretreatment,and temperature regime

Abies lasiocarpa (Hook.) Nutt. seeds germinated and the seedlings grew on under controlled environments (18 h photoperiod and 16/8 h thermoperiods; 10/10^oC, 20/20°C, 20/10°C, and 25/10°C). The experiment was concluded after 180 days, and the seedlings were scored for stem diameter and length. Four provenances were included: Colorado “high elevation”; (CO), Oregon 1 600 m a.s.l. (OR), British Columbia 1200 m a.s.l. (BC), and Finland (BCF; originally from British Columbia 1800 m a.s.l.). A cold‐moist pretreatment for 30 days improved germination and seedling growth in the BC and BCF seed lots, while the CO and OR seed lots were less affected. Daily fluctuating temperatures improved germination in the BC and BCF seed lots, the widest fluctuation 25/10°C producing the greatest germination. The average length of the seedlings was 34 mm at 20/20°C, 33 mm at 20/10°C, and 37 mm at 25/10°C; differences in hypocotyl length being most important. There were small but significant differences between the provenances in their reaction to temperature regime. The major part of the seedlings produced 2 or 3 flushes of growth in the 180 days period, the CO provenance having the most stabile buds and the OR provenance being the most labile.     

Growth,water relations,and survival of drought-exposed seedlings from six maternal families of honey mesquite (Prosopis glandulosa): responses to CO(2) enrichment

Low water availability is a leading contributor to mortality of woody seedlings on grasslands, including those of the invasive shrub Prosopis. Increasing atmospheric CO(2) concentration could favor some genotypes of this species over others if there exists intraspecific variation in the responsiveness of survivorship to CO(2). To investigate such variation, we studied effects of CO(2) enrichment on seedling survival in response to uniform rates of soil water depletion in six maternal families of honey mesquite (P. glandulosa Torr. var. glandulosa). Three families each from the arid and mesic extremes of the species' distribution in the southwestern United States were studied in environmentally controlled glasshouses. Relative water content at turgor loss and osmotic potential were not affected by CO(2) treatment. Increased atmospheric CO(2) concentration, however, increased growth, leaf production and area, and midday xylem pressure potential, and apparently reduced transpiration per unit leaf area of seedlings as soil dried. Consequently, CO(2) enrichment about doubled the fraction of seedlings that survived soil water depletion. Maternal families of honey mesquite differed in percentage survival of drought and in several other characteristics, but differences were of similar or of smaller magnitude compared with differences between CO(2) treatments. There was no evidence for genetic variation in the responsiveness of survivorship to CO(2). By increasing seedling survival of drought, increasing atmospheric CO(2) concentration could increase the abundance of honey mesquite where establishment is limited by water availability. Genetic types with superior ability to survive drought today, however, apparently will maintain that advantage in the future.     

Physiological and field growth responses of oak seedlings to warm storage

The effect of warm storage (15°C) for 0, 15 or 31 days, applied after cold storage until April, and date of lifting to cold storage on the physiological condition and field performance of two-year-old oak seedlings (Quercus robur L.) was investigated. Assessments before planting included plant moisture status, root and shoot dry weight, root growth potential (RGP), while after planting root growth, shoot growth phenology, shoot and root dry matter accumulation and stem quality were assessed. Warm storage effects were large, but lift date effects were small. Warm storage for 31 days reduced height and diameter growth, stem quality, total biomass, root growth, and reduced stem quality in the field, but 15 days storage had a smaller effect. Warm storage delayed bud break and shoot growth cessation but survival was unaffected. The depletion of food reserves during storage and low moisture availability might have caused shoot dieback leading to the development of poor quality stems. There was evidence that dry weight fraction of both the shoot tip and the taproot provided good information on the quality of the stock before planting. RGP was also a good indicator of quality. Electrolyte leakage readings from fine and taproots were not reliable indicators of plant quality.     

Effects of atmospheric CO(2) on longleaf pine: productivity and allocation as influenced by nitrogen and water

Longleaf pine (Pinus palustris Mill.) seedlings were exposed to two concentrations of atmospheric CO(2) (365 or 720 micro mol mol(-1)) in combination with two N treatments (40 or 400 kg N ha(-1) year(-1)) and two irrigation treatments (target values of -0.5 or -1.5 MPa xylem pressure potential) in open-top chambers from March 1993 through November 1994. Irrigation treatments were imposed after seedling establishment (i.e., 19 weeks after planting). Seedlings were harvested at 4, 8, 12, and 20 months. Elevated CO(2) increased biomass production only in the high-N treatment, and the relative growth enhancement was greater for the root system than for the shoot system. In water-stressed trees, elevated CO(2) increased root biomass only at the final harvest. Root:shoot ratios were usually increased by both the elevated CO(2) and low-N treatments. In the elevated CO(2) treatment, water-stressed trees had a higher root:shoot ratio than well-watered trees as a result of a drought-induced increase in the proportion of plant biomass in roots. Well-watered seedlings consistently grew larger than water-stressed seedlings only in the high-N treatment. We conclude that available soil N was the controlling resource for the growth response to elevated CO(2) in this study. Although some growth enhancement was observed in water-stressed trees in the elevated CO(2) treatment, this response was contingent on available soil N.     

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.     

Heat damage in boxed white spruce (Picea glauca [Moench.] Voss) seedlings: Its pre-planting detection and effect on field performance

This study investigated the effects of holding 1+0 PSB313a white spruce (Picea glauca (Moench.) Voss) seedlings in storage boxes at air temperatures of 5, 10, 20, 30 and 40°C for 12, 24, 48, 72 and 96 h before planting. The ability to detect physiological damage to seedlings as a result of such treatment, before planting, was also examined. After one growing season, no needle damage or mortality >8% was found for temperature treatments up to 20°C for 4 days. At 30°C and above, seedling damage and mortality increased, while bud flush, shoot height, stem diameter and shoot dry weight decreased with increasing temperature and duration of treatment. Seedling mortality in the field was 100% after the 40°C treatment exposure for 72 h or longer. Pre-planting needle electrolyte leakage was indicative of visible needle damage 14 days after planting, whereas stem electrolyte leakage and root growth potential were more closely related to end of season plantation mortality. Despite the lack of damage observed at 20°C or below, preplanting exposure of white spruce seedlings to temperatures above 5°C, during transportation and field storage, is not recommended.     

Effects of season,needle age and elevated atmospheric CO(2) on photosynthesis in Scots pine (Pinus sylvestris)

Five-year-old Scots pine (Pinus sylvestris L.) seedlings were grown in open-top chambers at ambient and elevated (ambient + 400 &mgr;mol mol(-1)) CO(2) concentrations. Net photosynthesis (A), specific leaf area (SLA) and concentrations of nitrogen (N), carbon (C), soluble sugars, starch and chlorophyll were measured in current-year and 1-year-old needles during the second year of CO(2) enrichment. The elevated CO(2) treatment stimulated photosynthetic rates when measured at the growth CO(2) concentration, but decreased photosynthetic capacity compared with the ambient CO(2) treatment. Acclimation to elevated CO(2) involved decreases in carboxylation efficiency and RuBP regeneration capacity. Compared with the ambient CO(2) treatment, elevated CO(2) reduced light-saturated photosynthesis (when measured at 350 &mgr;mol mol(-1) in both treatments) by 18 and 23% (averaged over the growing season) in current-year and 1-year-old needles, respectively. We observed significant interactive effects of CO(2) treatment, needle age and time during the growing season on photosynthesis. Large seasonal variations in photosynthetic parameters were attributed to changes in needle chemistry, needle structure and feedbacks governed by whole-plant growth dynamics. Down-regulation of photosynthesis was probably a result of reduced N concentration on an area basis, although a downward shift in the relationship between photosynthetic parameters and N was also observed.     

Influence of foliar N on foliar soluble sugars and starch of red spruce saplings exposed to ambient and elevated ozone

Red spruce (Picea rubens Sarg.) trees growing at high elevation in the northeastern United States have experienced decline in recent years but seedlings have proved to be relatively tolerant of a wide range of environmental stresses in controlled studies. One possible reason for the wide tolerance to stress in seedlings is their inherently large pool of carbohydrate reserves, which is available for maintenance during and regrowth after periods of stress. We tested for the effects of foliar N and exposure to ozone on foliar carbohydrate reserves of 20-year-old naturally regenerated saplings. The trees were maintained in native soil in 360-l containers for 5 years before the experiment. The year before the experiment, trees were fertilized with N,P,K to provide a population of trees from N deficient to N sufficient. As foliar N decreased below 0.9%, length of current-year shoots and specific needle area of current-year needles declined. Foliar N concentration was correlated with foliar sugar and starch concentrations, but relationships varied with time of year. Before bud break, foliar carbohydrates and N, in general, were positively correlated, and date of bud break was delayed in N-deficient trees. During active growth, foliar soluble sugars and N were positively correlated, but starch concentrations were negatively correlated with N. By late September, neither starch nor sugar concentration was correlated with N concentration. Ozone and foliar N concentrations did not interact to change foliar carbohydrate concentrations or shoot and needle growth in this relatively short-term study.