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PubMed1999

Fine root respiration in mature eastern white pine (Pinus strobus) in situ: the importance of CO(2) in controlled environments.

We measured seasonal fine root respiration rate in situ while controlling chamber temperature and [CO(2)]. Atmospheric [CO(2)] ([CO(2)](a)) and measured soil [CO(2)] ([CO(2)](s)) were alternately delivered to a cuvette containing intact fine roots of eastern white pine (Pinus strobus L.). Respiration rates were consistently higher in [CO(2)](a) than in [CO(2)](s) and were almost three times higher during midsummer. Respiration rates were immediately reversed after returning to the alternate [CO(2)] (i.e., [CO(2)](a) --> [CO(2)](s) --> [CO(2)](a), and vice versa) suggesting a direct effect of elevated [CO(2)] on apparent respiration. Soil-[CO(2)]-based respiration rates decreased with increasing [CO(2)] on a dry mass and tissue [N] basis. We conclude that estimates of soil CO(2) flux and soil carbon budgets may be improved by more completely accounting for the rhizosphere microclimate (i.e., soil temperature and [CO(2)](s)) during measurement of fine root respiration.

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PubMed1999

Acclimation of loblolly pine (Pinus taeda) seedlings to high temperatures.

To determine the extent to which loblolly pine seedlings (Pinus taeda L.) acclimate to high temperatures, seedlings from three provenances-southeastern Texas (mean annual temperature 20.3 degrees C), southwestern Arkansas (mean annual temperature 16.2 degrees C) and Chesapeake, Maryland (mean annual temperature 12.8 degrees C)-were grown at constant temperatures of 25, 30, 35 or 40 degrees C in growth chambers. After two months, only 14% of the seedlings in the 40 degrees C treatment survived, so the treatment was dropped from the experiment. Provenance and family differences were not significant for most measured variables. Total biomass was similar in the 25 and 30 degrees C treatments, and less in the 35 degrees C treatment. Foliage biomass was higher, and root biomass lower, in the 30 degrees C treatment compared with the 25 degrees C treatment. Net photosynthesis and dark respiration of all seedlings were measured at 25, 30 and 35 degrees C. Both net photosynthesis and dark respiration exhibited acclimation to the temperature at which the seedlings were grown. For each temperature treatment, the highest rate of net photosynthesis was measured at the growth temperature. Dark respiration rates increased with increasing measurement temperature, but the basal rate of respiration, measured at 25 degrees C, decreased from 0.617 &mgr;mol m(-2) s(-1) in the 25 degrees C treatment to 0.348 &mgr;mol m(-2) s(-1) in the 35 degrees C treatment, resulting in less carbon loss in the higher temperature treatments than if the seedlings had not acclimated to the growth conditions. Temperature acclimation, particularly of dark respiration, may explain why total biomass of seedlings grown at 30 degrees C was similar to that of seedlings grown at 25 degrees C.

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PubMed1999

Magnesium nutrition and photosynthesis in Pinus radiata: clonal variation and influence of potassium.

Magnesium (Mg) nutrition and photosynthesis were studied in clones of Pinus radiata D. Don grown in sand culture for 21 weeks at four Mg concentrations (0.008, 0.04, 0.2 and 0.4 mM) and three potassium (K) concentrations (0.25, 0.5 and 2.5 mM). We found significant clonal variation in Mg nutrition of P. radiata. Plants grown at 0.04 mM [Mg] or less showed pronounced visible symptoms of foliar Mg deficiency. Net photosynthetic rate and leaf conductance were closely related to shoot Mg concentrations below a concentration of 0.6 mg Mg g(DW) (-1). Potassium enhanced the development of visible symptoms of foliar Mg deficiency. At the lowest Mg concentration tested (0.008 mM), the severity of needle chlorosis and necrosis increased with increasing K concentration in the culture solution. With increasing Mg concentration, 2.5 mM [K] in the culture solution markedly increased root Mg concentration, but decreased shoot Mg concentration, suggesting that excessive K inhibited Mg mobilization from roots to shoots. Rates of growth and photosynthesis were both severely inhibited at 0.008 mM [Mg].

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PubMed1999

Effects of water status and soil fertility on the C-isotope signature in Pinus radiata.

The efficiency with which trees use water is a major determinant of growth under water-limited conditions. We investigated whether increased access to water and nutrients alters water-use efficiency in Pinus radiata D. Don. Intrinsic transpiration efficiency, defined here as the ratio of CO(2) assimilated and water transpired at a given vapor pressure deficit, is determined by the difference between ambient atmospheric CO(2) concentration (c(a)) and leaf intercellular CO(2) concentration (c(i)). The mean value of c(i)/c(a) can be inferred from an analysis of carbon isotope discrimination (Delta) in wood samples. A total of 117 trees, growing at sites with widely varying soil and climatic conditions in Australia and New Zealand, were cored and distinct annual rings were analyzed for their carbon isotope ratio, and correlated with rainfall during the July-June growing season in the year in which the wood was grown. Where possible, carbon isotope ratios were compared for different years within the same trees. The c(i)/c(a) ratio decreased with decreasing water availability, suggesting that intrinsic transpiration efficiency increased with decreasing water availability. An increase in growing season rainfall of 900 mm resulted in an increase in Delta of about 2.0 per thousand, corresponding to a decrease in intrinsic transpiration efficiency of approximately 24%. A stronger relationship was obtained when carbon isotope discrimination was expressed as a function of the ratio of rainfall to potential transpiration. Carbon isotope discrimination was also negatively correlated with mean annual vapor pressure deficit at different sites. In contrast, nutrient availability had no significant effect on carbon isotope discrimination.

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PubMed1999

Diurnal changes in water conduction in loblolly pine (Pinus taeda) and Virginia pine (P. virginiana) during soil dehydration.

We studied diurnal changes in water conduction during soil dehydration in 37-month-old seedlings of one Virginia pine (Pinus virginiana Mill.) and two loblolly pine (P. taeda L.) sources, one from North Carolina (NC) and the other from the "Lost Pines" areas of Texas (TX), in an environmentally controlled growth chamber. For seedlings of similar biomass, the TX source had higher values of transpiration, needle conductance, and plant hydraulic conductivity under well-watered conditions than the NC source. Under dry soil conditions, the TX source had lower values of water conduction than the NC source. The Virginia pine source responded similarly to the TX source under both well-watered and dry soil conditions. For all three pine sources, gradients between soil and needle water potentials were greatest when the seedlings were moderately stressed. The TX and Virginia pine sources had higher gradients and lower daytime needle water potentials under moderate stress conditions than the NC source. Predawn needle water potentials did not differ among the pine sources. We conclude that the TX and Virginia pine sources have decreased daytime needle water potentials and increased water potential gradients during the daytime under moderate stress conditions, but with no disruption of recovery at predawn. The greater rates of transpiration and water conduction by the TX source compared with the NC source under well-watered conditions suggest a means by which growth can be maximized prior to the onset of drought, thereby enhancing survival of loblolly pines in drought-prone environments.

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PubMed1986

Stomatal and nonstomatal limitations to net photosynthesis in Pinus taeda L. under different environmental conditions.

Net photosynthesis, transpiration and stomatal conductance of two-year-old Pinus taeda L. seedlings were compared under various environmental conditions. Responses to air temperature, irradiance, ambient CO(2) concentration, absolute humidity deficit and xylem pressure potential were examined. The seedlings exhibited little response to a wide range of absolute humidity deficits (7 to 16 g m(-3)) and temperatures (20 to 35 degrees C), but were sensitive to changes in water deficit, irradiance and CO(2) concentration. Net photosynthesis and stomatal conductance were linearly related under all of the environmental conditions measured. However, the gas phase limitation to photosynthesis was generally small (20 to 30%). It was concluded that although the stomatal response was closely coupled to changes in photosynthesis, internal limitations, rather than the rate of gaseous diffusion of CO(2), were primarily responsible for limiting photosynthesis.

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PubMed1986

Growth and physiological responses of Pinus ponderosa Dougl ex P. Laws. to long-term elevated CO(2) concentrations.

Seven-year-old ponderosa pine (Pinus ponderosa Dougl. ex P. Laws.) saplings and one- and two-year-old ponderosa pine seedlings of a Sierra Nevada and a Rocky Mountain seed source, respectively, were exposed to CO(2)-enriched atmospheres in an outdoor open-top chamber facility for 2.5 years. Seedling growth (main stem diameter, height, volume) increased with increasing CO(2) concentration, though the two populations exhibited different patterns of response. By the beginning of the last growth season, however, the trees under the highest CO(2) concentrations showed signs of stress that included accelerated needle abscision, chlorosis, and apparent alteration of tolerance to heat. The stress response is at least partly attributable to elevated foliar temperatures resulting from CO(2)-induced stomatal closure, which in turn lowered transpirational cooling of needles.

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PubMed1986

Evidence of competition for photosynthates between growth processes and oleoresin synthesis in Pinus taeda L.

A group of 10 pines (Pinus taeda L.) in a 35-year-old stand in Central Louisiana was sampled periodically from March through December 1984 for oleoresin flow from small wounds. Oleoresin yield was closely associated with periods of calculated moderate soil water deficit and presumed growth reduction. The period of maximum oleoresin flow coincided with the period of calculated maximum soil water depletion. Calculated potential evapotranspiration reached a maximum in late June, but water deficits did not reach maximum levels until September. Apparently, moderate seasonal water deficits that limit growth, but do not limit photosynthesis and translocation of photosynthates, favor differentiation processes, such as oleoresin synthesis. Because oleoresin is a primary defense against southern pine beetle (Dendroctonus frontalis Zimm.) attack in pines, recognition of these relationships should contribute to an understanding of southern pine beetle-host tree interactions.

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PubMed1986

Accumulation of starch in roots and foliage of loblolly pine (Pinus taeda L.): effects of season, site and fertilization.

Accurate predictions of stand response to nitrogen fertilization require a better understanding of the plant processes associated with stand nutrition. This study examined the relationship between stand nutrition and starch accumulation in roots and foliage of loblolly pine (Pinus taeda L.). A field study was located on the lower Coastal Plain of North Carolina on four sites of differing soil types. Stands of 8-year-old trees were fertilized with nitrogen (225 kg ha(-1)) or nitrogen plus phosphorus (225 kg N ha(-1) + 75 kg P ha(-1)) in 1981. Roots and foliage were collected from five trees per plot at six sampling dates from July 1983 through January 1985 and analyzed for starch. Starch concentrations showed seasonal fluctuations, with levels generally highest in early spring prior to budbreak. Root starch concentrations were lowest in autumn, whereas foliar concentrations were lowest in winter.

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PubMed1986

Relationships between nutrient status, carbon assimilation and water use efficiency in Pinus radiata (D. Don) needles.

The productivity of Pinus radiata D. Don plantations can be increased on many soils by N and P fertilization. Foliar analyses are usually taken as a guide for nutrient management, but there is little information about the relationship of foliar nutrient status to foliar physiology. This paper provides such information from N and P fertilization experiments. The concentrations and contents of N and P in the needles of P. radiata change because of accumulation during needle growth, retranslocation during shoot growth, and replenishment from current uptake during periods of little shoot growth. Up to 60% P and 50% N may be retranslocated in a single growing season from needles less than one year old. Three years after fertilizer application to a 10-year-old thinned P. radiata stand, net carbon assimilation rate per unit leaf area was in the order NP>P>N. There were positive associations between foliar P and assimilation at full sunlight, diffusive conductance and quantum yield. Foliar N was negatively associated with these variables. Higher concentrations of P and N were both accompanied by higher instantaneous water use efficiency by the foliage. Estimated trends in whole tree assimilation were similar to those in aboveground biomass accumulation. The interplay between the dynamic nature of nutrients in the foliage and their influence on the physiology of needles is discussed.

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PubMed2000

Water availability and branch length determine delta(13)C in foliage of Pinus pinaster.

The stable carbon isotope composition (delta(13)C) of foliage integrates signals resulting from environmental and hydraulic constraints on water movement and photosynthesis. We used branch length as a simple predictor of hydraulic constraints to water fluxes and determined the response of delta(13)C to varying water availability. Foliage up to 6 years old was taken from Pinus pinaster Ait. trees growing at four sites differing in precipitation (P; 414-984 mm year(-1)) and potential evaporation (ET; 1091-1750 mm year(-1)). Branch length was the principal determinant of temporal trends in delta(13)C. The strong relationship between delta(13)C and branch length was a function of hydraulic conductance, which was negatively correlated with branch length (r(2) = 0.84). Variation in P and ET among sites was reflected in delta(13)C, which was negatively correlated with P/ET (r(2) = 0.66). However, this analysis was confounded by differences in branch length. If the effects of branch length on delta(13)C were first removed, then the 'residual' delta(13)C was more closely related to P/ET (r(2) = 0.99), highlighting the importance of accounting for variation in hydraulic constraints to water flux between sites and years. For plant species that exhibit considerable phenotypic plasticity in response to changes in environment (e.g., variation in leaf area, branch length and number, or stem form), the environmental effects on delta(13)C in foliage can only be reliably assessed if deconvoluted from hydraulic constraints.

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PubMed2000

Effect of pruning on nitrogen dynamics within crowns of Pinus radiata.

Stem injection of (15)N-labeled ammonium sulfate was used to determine effects of pruning on canopy nitrogen dynamics in open-grown Pinus radiata D. Don in New Zealand. Trees were planted in July 1990 and the isotope introduced in December 1994. Tree crowns were divided into three zones: base section, from which branches of pruned trees were removed; mid section, between the pruned zone and the height of the trees at the start of the year in which they were pruned; and top section, which grew predominantly after the isotope was applied. Pruning removed 32% of the green crown length, representing 75% of foliage biomass. Needles were sampled from each region of the crown until July 1996. Branch growth was used to predict foliage biomass for each sampling occasion. Approximately 45% of the applied isotope was recovered from needles sampled in December 1994 (1 week after application and immediately before pruning), two-thirds of which occurred in needles in the base section. Thereafter, changes in isotope content of needles in the base section of unpruned trees largely reflected foliage biomass fluctuations and dilution of the isotope by continued uptake from the unlabeled soil nitrogen pool. Recovery of isotope in needles from the mid-crown section increased by 58 and 86% from December 1994 to July 1995 in control and pruned trees, respectively. Within this crown section, there was evidence of isotope translocation from old to new needles, with both isotope dilution and efflux observed in the needle cohorts that had been present at the time the isotope was applied. Therefore, isotope dynamics did not reflect the dynamics of the total nitrogen pool in the mid-crown section. By July 1996, a small proportion of the applied isotope was recovered from the new foliage formed in the top section of the crown. Within the top section, isotope dynamics closely matched total nitrogen fluxes. Pruning the lower crown did not affect nitrogen dynamics elsewhere in the crown for the following 18 months.

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PubMed2000

Genetic and environmental control of seasonal carbohydrate dynamics in trees of diverse Pinus sylvestris populations.

We explored environmental and genetic factors affecting seasonal dynamics of starch and soluble nonstructural carbohydrates in needle and twig cohorts and roots of Scots pine (Pinus sylvestris L.) trees of six populations originating between 49 degrees and 60 degrees N, and grown under common garden conditions in western Poland. Trees of each population were sampled once or twice per month over a 3-year period from age 15 to 17 years. Based on similarity in starch concentration patterns in needles, two distinct groups of populations were identified; one comprised northern populations from Sweden and Russia (59-60 degrees N), and another comprised central European populations from Latvia, Poland, Germany and France (49-56 degrees N). Needle starch concentrations of northern populations started to decline in late spring and reached minimum values earlier than those of central populations. For all populations, starch accumulation in spring started when minimum air temperature permanently exceeded 0 degrees C. Starch accumulation peaked before bud break and was highest in 1-year-old needles, averaging 9-13% of dry mass. Soluble carbohydrate concentrations were lowest in spring and summer and highest in autumn and winter. There were no differences among populations in seasonal pattern of soluble carbohydrate concentrations. Averaged across all populations, needle soluble carbohydrate concentrations increased from about 4% of needle dry mass in developing current-year needles, to about 9% in 1- and 2-year-old needles. Root carbohydrate concentration exhibited a bimodal pattern with peaks in spring and autumn. Northern populations had higher concentrations of fine-root starch in spring and autumn than central populations. Late-summer carbohydrate accumulation in roots started only after depletion of starch in needles and woody shoots. We conclude that Scots pine carbohydrate dynamics depend partially on inherited properties that are probably related to phenology of root and shoot growth.

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PubMed2000

Effects of serpentine soil factors on Virginia pine (Pinus virginiana) seedlings.

Effects of simulated serpentine soil conditions (elevated Mg:Ca ratio and Ni concentration) on seedlings from populations of Virginia pine (Pinus virginiana Mill.) from serpentine and non-serpentine sites were evaluated in sand culture. We determined (1) how seedlings are affected by elevated Mg:Ca ratio and Ni concentrations, (2) if there are interactive effects between Mg:Ca ratio and Ni concentrations on seedling growth, needle pigment concentrations, and nutrition, and (3) if Virginia pine populations from serpentine areas are edaphic ecotypes. A Mg:Ca ratio of 5 and 50 microM Ni both reduced seedling growth compared with control seedlings grown in the presence of the standard Mg:Ca ratio of 0.5 and no Ni. Interactive effects between Mg:Ca ratio and Ni concentrations were highly significant for growth, foliar pigments, and needle and root elemental concentrations. Nickel-mediated reductions in growth and foliar pigment concentrations were less at the serpentine Mg:Ca ratio of 5 than at the standard (non-serpentine) Mg:Ca ratio of 0.5. Foliar N was reduced by Ni concentrations as low as 10 microM, and foliar and root K, Ca and P concentrations were significantly reduced by Ni concentrations above 25 microM, with greater reductions at a Mg:Ca ratio of 0.5 than at a Mg:Ca ratio of 5. There were no population x serpentine soil factor interactions for seedling growth, foliar pigment concentrations, or nutrition, suggesting that seedlings from trees growing on serpentine soils are not edaphic ecotypes. We conclude that serpentine conditions present at the site of seed collection have not resulted in the selection of edaphic ecotypes of Virginia pine with respect to Mg:Ca ratio and Ni concentration.

Asbestos, Serpentine↗

PubMed2000

Frost hardening of Pinus radiata seedlings: effects of temperature on relative growth rate, carbon balance and carbohydrate concentration.

Pinus radiata (D. Don) seedlings were grown for 100 days at day/night temperatures of 10/1, 15/1, 20/1 and 25/1 degrees C, to determine whether temperatures above a threshold of 5 degrees C influence frost hardiness development. Relationships between hardening and relative growth rate, carbohydrate concentration and net carbon balance were also investigated. Seedlings hardened at a nearly constant rate in each treatment, although the rate of hardening was strongly temperature dependent. It increased as the temperature declined, but in a curvilinear fashion. Temperatures below 9.5 degrees C were effective in hardening the seedlings. During the daily temperature cycle, dehardening occurred at temperatures above the threshold, whereas hardening occurred at temperatures below the threshold. The net difference between the two processes determined the development of frost hardiness. The development of frost hardiness was negatively correlated with relative growth rate and positively correlated with the accumulation of starch and sugars. We conclude that frost hardening is a complex process that is causally linked to carbohydrate concentrations.

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PubMed2000

Dynamics of symbiotic establishment between an IAA-overproducing mutant of the ectomycorrhizal fungus Hebeloma cylindrosporum and Pinus pinaster.

To clarify the early steps of symbiotic establishment, we studied the dynamics of Pinus pinaster (Ait.) Sol. tap root colonization and mycorrhiza formation by an IAA-overproducing mutant of the ectomycorrhizal fungus Hebeloma cylindrosporum Romagnesi and by the corresponding wild type strain. Differences between wild type and mutant strains were quantitative rather than qualitative and were detected two days after inoculation. Both fungal strains established a typical Hartig net when they colonized the tap roots. Consequently, colonized tap roots exhibited features of a true mycorrhiza and fungal colonization enhanced plant growth. Fungal colonization and Hartig net formation were more rapid with the mutant than with the wild type. Colonization, especially with the mutant strain, increased rhizogenesis and the production of mycorrhizas. The mutant formed a hypertrophic Hartig net in tap roots and mycorrhizal short roots and we obtained evidence that the process of short root transformation into mycorrhiza started before their emergence from the tap root. Hyphae of the Hartig net in the tap root penetrated the cortex of young lateral roots at the beginning of their elongation, after the endodermis layer broke under the pressure of the elongating lateral root. Colonization was inhibited when triiodobenzoic acid was added to the culture medium, providing circumstantial evidence that auxin is involved in mycorrhiza formation.

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PubMed2000

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.

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PubMed2000

Seasonal CO(2) assimilation and stomatal limitations in a Pinus taeda canopy.

Net CO(2) assimilation (A(net)) of canopy leaves is the principal process governing carbon storage from the atmosphere in forests, but it has rarely been measured over multiple seasons and multiple years. I measured midday A(net) in the upper canopy of maturing loblolly pine (Pinus taeda L.) trees in the piedmont region of the southeastern USA on 146 sunny days over 36 months. Concurrent data for leaf conductance and photosynthetic CO(2) response curves (A(net)-C(i) curves) were used to estimate the relative importance of stomatal limitations to CO(2) assimilation in the field. In fully expanded current-year and 1-year-old needles, midday light-saturated A(net) was constant over much of the growing season (5-6 &mgr;mol CO(2) m(-2) s(-1)), except during drought periods. During the winter season (November-March), midday A(net) of overwintering needles varied in proportion to leaf temperature. Net CO(2) assimilation at light saturation occurred when daytime air temperatures exceeded 5-6 degrees C, as happened on more than 90% of the sunny winter days. In both age classes of foliage, winter carbon assimilation accounted for approximately 15% of the daily carbon assimilation on sunny days throughout the year, and was relatively insensitive to year-to-year differences in temperature during this season. However, strong stomatal limitations to A(net) occurred as a result of water stress associated with freezing cycles in winter. During the growing season, drought-induced water stress produced the largest year-to-year differences in seasonal CO(2) assimilation on sunny days. Seasonal A(net) was more drought sensitive in current-year needles than in 1-year-old needles. Relative stomatal limitations to daily integrated A(net) were approximately 40% over the growing season, and summer drought rather than high temperatures had the largest impact on summer A(net) and integrated annual CO(2) uptake in the upper crown. Despite significant stomatal limitations, a long duration of near-peak A(net) in the upper crown, particularly in 1-year-old needles, conferred high seasonal and annual carbon gain.

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