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Effects of exogenous diamines on the interaction between ectomycorrhizal fungi and adventitious root formation in Scots pine in vitro.

Production of free and conjugated polyamines by two ectomycorrhizal fungi, Pisolithus tinctorius (Pers.) Coker and Couch and Paxillus involutus (Batsch) Fr., was studied in vitro. Spermidine was the main polyamine in the mycelium of both fungi. Paxillus involutus also produced large amounts of the diamine putrescine, whereas Pisolithus tinctorius contained traces of the diamine cadaverine and released into the culture medium an unknown compound probably related to cadaverine or N-methylputrescine. Both fungi accelerated adventitious root formation and increased subsequent root growth of Scots pine (Pinus sylvestris L.) hypocotyl cuttings in vitro. Exogenous cadaverine enhanced rooting caused by Pisolithus tinctorius and also promoted mycorrhiza formation by the fungus. Putrescine and Paxillus involutus had a synergistic effect on root initiation, but not on subsequent root growth. We conclude that specific diamines may be involved in the interaction between ectomycorrhizal fungi and adventitious root formation in Scots pine, and that the effects of specific exogenous polyamines are dependent on the fungal strain and its ability to produce these compounds. The finding that Paxillus involutus enhanced rooting and root growth without mycorrhiza formation indicates that fungal-induced rooting is not necessarily related to visible mycorrhiza formation.

Basidiomycota↗

Comparing the influence of site quality, stand age, fire and climate on aboveground tree production in Siberian Scots pine forests.

Temporal patterns of stem and needle production and total aboveground net primary production (ANPP) were studied at the tree and stand level along four chronosequences of Siberian Scots pine (Pinus sylvestris L.) forests differing in site quality (poor lichen type or the more fertile Vaccinium type) and in frequency of surface fires (unburned, moderately burned (fire return interval of approximately 40 years), or heavily burned (fire return interval of approximately 25 years)). The maximum range of variability in aboveground production was quantified for: (1) possible long-term changes in site quality; (2) stand age; (3) non-stand-replacing, recurring surface fires; and (4) interannual climate variability. For (1) and (2), total ANPP was low in the lichen-type chronosequence, reached a maximum of 170 g C m(-2) year(-1) after 100 years and decreased to 100 g C m(-2) year(-1) in older stands. Maximum ANPP in the Vaccinium-type chronosequence was 340 g C m(-2) year(-1) and occurred earlier in the 53-year-old stand than in the other stands. Along the lichen-type chronosequences, peak ANPP was paralleled by maximum carbon allocation to stem growth. (3) In mature trees, damage by recurrent surface fires decreased stem growth by 17 +/- 19% over a 10-year period relative to pre-fire values. At longer timescales, ANPP was hardly affected by fire-related differences in mortality. (4) Needle- plus stem-NPP, reconstructed for a 3-year period, varied within a range of 15 g C m(-2) year(-1) in the lichen-type stands and 35 g C m(-2) year(-1) in the Vaccinium-type stands. For the same period, the coefficient of variance was higher for needle-NPP (20 +/- 10%) than for stem-NPP (12 +/- 7%). Needle- and stem-NPP did not covary in time. Most 30-year time series of stem-NPP at the tree level exhibited strong autocorrelation. In older trees, stem-NPP was positively correlated with growing season precipitation. Thus, the factors driving variability in ANPP ranked according to their maximum influence as: stand age (controlled by the frequency of stand-replacing fires) > site quality > growth depression because of surface fire damage approximately equal age-related reduction in ANPP > interannual variability approximately equal long-term effects of fire (stand density reduction). In lichen-type forests, we found that ANPP at the landscape level declined sharply when the interval between stand-replacing fires was less than 120 years, illustrating that fire strongly influences ANPP of boreal Scots pine forests.

Climate↗

A multichannel automated chamber system for continuous measurement of forest soil CO2 efflux.

We developed a fast-response multi-chamber system for measuring soil-surface CO2 efflux (Fc). The chambers (90 x 90 x 50 cm, L x W x H) had lids that opened and closed automatically, and were connected in parallel to a single CO2 analyzer equipped with a 16-channel gas sampler. Between measurements the chamber lids were raised to allow precipitation and leaf litter to reach the enclosed soil surface. When a chamber was closed, it was ventilated with well-buffered ambient air (125 l min-1) that entered by an inlet on one chamber sidewall and exited through a large vent on the opposite sidewall. The pressure difference between the inside and outside of the chamber was less than 0.22 Pa. Two additional mixing fans maintained an air speed of 0.3 +/- 0.1 m s-1 at 20 cm above the soil surface. Air was withdrawn continuously from the inlets and outlets of each chamber, and fed sequentially to an infrared CO2 analyzer. With this system, we measured Fc in a 40-year-old temperate Pinus densiflora Sieb. & Zucc. forest from February 8 to May 30, 2001. Mean Fc increased steadily from 0.9 micro mol m-2 s-1 at the beginning of February to 4.6 micro mol m-2 s-1 by the end of May. There was a statistically significant correlation between Fc and surface soil temperature (r = 0.896; P < 0.0001), and the Q10 value was 2.8. Spatial variation of Fc was higher in the non-growing season than in the growing season. Measurements were not interrupted by either rain or snow.

Botany↗

Stem sapwood permeability in relation to crown dominance and site quality in self-thinning fire-origin lodgepole pine stands.

Stem sapwood hydraulic permeability, tree leaf area, sapwood basal area, earlywood to latewood ratio of annual rings, radial variation in hydraulic permeability and stem hydraulic capacity were examined in dominant (D), codominant (CD) and suppressed (SP) lodgepole pine (Pinus contorta Dougl. ex Loud.) trees growing on medium and poor sites. Hydraulic permeability on a sapwood area basis (ks) was lower in suppressed trees (0.71 x 10(-12) m2) compared to dominants (1.97 x 10(-12) m2) and codominants (1.79 x 10(-12) m2), and higher on medium than on poor sites. The leaf/sapwood area ratio (S) varied with crown dominance position (D > CD > SP) but not by site type. Leaf specific conductivity (kL) did not vary between crown classes or site types. The relationship between leaf area and stem hydraulic supply capacity (Q*) was strong, but differed among crown classes. Dominant trees and trees from the medium sites had a greater proportion of earlywood in outer rings of sapwood than suppressed trees. Sapwood permeability declined from the cambium to the sapwood-heartwood boundary in all samples, but the decline was more gradual in dominant trees compared to codominant and suppressed trees; differences in the radial variation in sapwood permeability may be related to differences in S. Sapwood permeability is positively related to crown dominance, whereas subdominant (CD and SP) trees have greater Q* in relation to leaf area, leading us to propose that this may give subdominant trees a survival advantage, slowing self-thinning.

Fires↗

Variation in morphological and biochemical O3 injury attributes of mature Jeffrey pine within canopies and between microsites.

Crown morphology and leaf tissue chemical and biochemical attributes associated with ozone (O3) injury were assessed in the lower, mid- and upper canopy of Jeffrey pine (Pinus jeffreyi Grev. & Balf.) growing in mesic and xeric microsites in Sequoia National Park, California. Microsites were designated mesic or xeric based on topography and bole growth in response to years of above-average precipitation. In mesic microsites, canopy response to O3 was characterized by thinner branches, earlier needle fall, less chlorotic leaf mottling, and lower foliar antioxidant capacity, especially of the aqueous fraction. In xeric microsites, canopy response to O3 was characterized by higher chlorotic leaf mottling, shorter needles, lower needle chlorophyll concentration, and greater foliar antioxidant capacity. Increased leaf chlorotic mottle in xeric microsites was related to drought stress and increased concurrent internal production of highly reactive oxygen species, and not necessarily to stomatal O3 uptake. Within-canopy position also influenced the expression of O3 injury in Jeffrey pine.

Ozone↗

Developmental lignification and seasonal variation in beta-glucosidase and peroxidase activities in xylem of Scots pine, Norway spruce and silver birch.

We examined the relationship between beta-glucosidase and peroxidase activities and xylem lignification in the stems of Scots pine (Pinus sylvestris L.), Norway spruce (Picea abies (L.) Karst.) and silver birch (Betula pendula Roth) during the 1999 growing season. Examination of stem cross sections stained with safranin and Alcian blue for lignin and cellulose, respectively, indicated that radial growth of pine and spruce xylem began in late May, whereas the growth of birch xylem was initiated 2 weeks later. Lignification began soon after thickening of the newly formed cell walls, i.e., upon deposition of cellulose. Hydrolysis of the synthetic beta-glucosidase substrate p-nitrophenyl-beta-O-D-glucopyranoside was correlated with radial growth and lignification in the xylem of both conifers, but the relationship between lignification and the hydrolysis of coniferin by beta-glucosidase was not obvious. Beta-glucosidase activities in the xylem of silver birch were low and did not correlate with growth or lignification with either substrate. An increase in peroxidase activity was detected at the initiation of growth and lignification in the conifers and during growth and lignification in silver birch, but high peroxidase activities were also measured outside the growth period during late autumn, winter and early spring.

Betula↗

Freezing tolerance of conifer seeds and germinants.

Survival after freezing was measured for seeds and germinants of four seedlots each of interior spruce (Picea glauca x engelmannii complex), lodgepole pine (Pinus contorta Dougl. ex Loud.), Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) and western red cedar (Thuja plicata Donn ex D. Donn). Effects of eight seed treatments on post-freezing survival of seeds and germinants were tested: dry, imbibed and stratified seed, and seed placed in a growth chamber for 2, 5, 10, 15, 20 or 30 days in a 16-h photoperiod and a 22/17 degrees C thermoperiod. Survival was related to the water content of seeds and germinants, germination rate and seedlot origin. After freezing for 3 h at -196 degrees C, dry seed of most seedlots of interior spruce, Douglas-fir and western red cedar had 84-96% germination, whereas lodgepole pine seedlots had 53-82% germination. Freezing tolerance declined significantly after imbibition in lodgepole pine, Douglas-fir and interior spruce seed (western red cedar was not tested), and mean LT50 of imbibed seed of these species was -30, -24.5 and -20 degrees C, respectively. Freezing tolerance continued to decline to a minimum LT50 of -4 to -7 degrees C after 10 days in a growth chamber for interior spruce, Douglas-fir and lodgepole pine, or after 15 days for western red cedar. Minimum freezing tolerance was reached at the stage of rapid hypocotyl elongation. In all species, a slight increase in freezing tolerance of germinants was observed once cotyledons emerged from the seed coat. The decrease in freezing tolerance during the transition from dry to germinating seed correlated with increases in seed water content. Changes in freezing tolerance between 10 and 30 days in the growth chamber were not correlated with seedling water content. Within a species, seedlots differed significantly in freezing tolerance after 2 or 5 days in the growth chamber. Because all seedlots of interior spruce and lodgepole pine germinated quickly, there was no correlation between seedlot hardiness and rate of germination. Germination rate and freezing tolerance of Douglas-fir and western red cedar seedlots was negatively correlated. There was a significant correlation between LT50 after 10 days in the growth chamber and minimum spring temperature at the location of seedlot origin for interior spruce and three seedlots of western red cedar, but no relationship was apparent for lodgepole pine and Douglas-fir.

Freezing↗

Interactive effects of temperature and precipitation on soil respiration in a temperate maritime pine forest.

Soil respiration (SR) was monitored periodically throughout 2001 in a Scots pine (Pinus sylvestris L.) stand located in the Belgian Campine region. As expected for a temperate maritime forest, temperature was the dominant control over SR during most of the year. However, during late spring and summer, when soil water content (SWC) was limiting, SR was insensitive to temperature (Q(10) = 1.24). We observed that during prolonged rain-free periods, when SWC was less than 15% (v/v), SR decreased dramatically (up to 50%) and SWC took over control of SR. During such drought periods, however, rain events sometimes stimulated SR and restored temperature control over SR, even though SWC in the mineral soil was low. We hypothesize that restoration of temperature control occurred only when rain events adequately rewetted the uppermost soil layers, where most of the respiratory activity occurred. To quantify the rewetting capacity of rain events, an index (I(w)) was designed that incorporated rainfall intensity, time elapsed since the last rain event, and atmospheric vapor pressure deficit (a proxy for evaporative water losses). To simulate SR fluxes, a model was developed that included the effects of soil temperature and, under drought and non-rewetting conditions (I(w) and SWC < threshold), an SWC response function. The model explained 95% of the temporal variability in SR observed during summer, whereas the temperature function alone explained only 73% of this variability. Our results revealed that, in addition to temperature and SWC, rain plays a role in determining the total amount of carbon released from soils, even in a maritime climate.

Cell Respiration↗

Contrasting effects of elevated carbon dioxide concentration and temperature on Rubisco activity, chlorophyll fluorescence, needle ultrastructure and secondary metabolites in conifer seedlings.

Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) seedlings were grown for 50 days in growth chambers in an ambient or twice ambient carbon dioxide concentration ([CO2]) at a day/night temperature of 19/12 degrees C or 23/16 degrees C. Although elevated [CO2] (EC) had only slight effects on the growth parameters measured, elevated temperature (ET) increased above ground dry mass of both species. Among treatments, biomass accumulation of both species was greatest in the combined EC + ET treatment. The EC treatment induced thylakoid swelling and increased numbers of plastoglobuli observed in Scots pine needles. Although EC had little effect on Rubisco protein or N concentration of needles, ET had a large effect on N-containing compounds and enhanced N allocation from 1-year-old needles. Terpenoids were more responsive to EC and ET than total phenolics. Generally, terpene concentrations were reduced by EC and increased by ET. Increased terpenoid concentrations in response to ET might be associated with thermotolerance of photosynthesis. In Norway spruce, EC decreased total phenolic concentrations in needles, probably as a result of increased growth. We conclude that, in seedlings of these boreal species, the effects of elevated [CO2] on the studied parameters were small compared with the effects of elevated temperature.

Carbon Dioxide↗

Reliance on stored water increases with tree size in three species in the Pacific Northwest.

In tall old forests, limitations to water transport may limit maximum tree height and reduce photosynthesis and carbon sequestration. We evaluated the degree to which tall trees could potentially compensate for hydraulic limitations to water transport by increased use of water stored in xylem. Using sap flux measurements in three tree species of the Pacific Northwest, we showed that reliance on stored water increases with tree size and estimated that use of stored water increases photosynthesis. For Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), water stored in xylem accounted for 20 to 25% of total daily water use in 60-m trees, whereas stored water comprised 7% of daily water use in 15-m trees. For Oregon white oak (Quercus garryana Dougl. ex Hook.), water stored in xylem accounted for 10 to 23% of total daily water use in 25-m trees, whereas stored water comprised 9 to 13% of daily water use in 10-m trees. For ponderosa pine (Pinus ponderosa Dougl. ex Laws.), water stored in xylem accounted for 4 to 20% of total daily water use in 36-m trees, whereas stored water comprised 2 to 4% of daily water use in 12-m trees. In 60-m Douglas-fir trees, we estimated that use of stored water supported 18% more photosynthesis on a daily basis than would occur if no stored water were used, whereas 15-m Douglas-fir trees gained 10% greater daily photosynthesis from use of stored water. We conclude that water storage plays a significant role in the water and carbon economy of tall trees and old forests.

Northwestern United States↗

Field measurements of root respiration indicate little to no seasonal temperature acclimation for sugar maple and red pine.

Increasing global temperatures could potentially cause large increases in root respiration and associated soil CO2 efflux. However, if root respiration acclimates to higher temperatures, increases in soil CO2 efflux from this source would be much less. Throughout the snow-free season, we measured fine root respiration in the field at ambient soil temperature in a sugar maple (Acer saccharum Marsh.) forest and a red pine (Pinus resinosa Ait.) plantation in Michigan. The objectives were to determine effects of soil temperature, soil water availability and experimental N additions on root respiration rates, and to test for temperature acclimation in response to seasonal changes in soil temperature. Soil temperature and soil water availability were important predictors of root respiration and together explained 76% of the variation in root respiration rates in the red pine plantation and 71% of the variation in the sugar maple forest. Root N concentration explained an additional 6% of the variation in the sugar maple trees. Experimental N additions did not affect root respiration rates at either site. From April to November, root respiration rates measured in the field increased exponentially with increasing soil temperature. For sugar maple, long-term Q10 values calculated from the field data were slightly, but not significantly, less than short-term Q10 values determined for instantaneous temperature series conducted in the laboratory (2.4 versus 2.62.7). For red pine, long-term and short-term Q10 values were similar (3.0 versus 3.0). Sugar maple root respiration rates at constant reference temperatures of 6, 18 and 24 degrees C were measured in the laboratory at various times during the year when field soil temperatures varied from 0.4 to 16.8 degrees C. No relationship existed between ambient soil temperature just before sampling and root respiration rates at 6 and 18 degrees C (P = 0.37 and 0.86, respectively), and only a very weak relationship was found between ambient soil temperature and root respiration at 24 degrees C (P = 0.08, slope = 0.09). We conclude that root respiration in these species undergoes little, if any, acclimation to seasonal changes in soil temperature.

Acclimatization↗

Methyl jasmonate induces changes mimicking anatomical defenses in diverse members of the Pinaceae.

Conifers have defenses such as the production of phenolic compounds and resins that can be induced by bark beetles and other invading organisms, but the signaling agents involved are unknown. The anatomical effects of methyl jasmonate (MJ), a potent inducer of certain plant defenses, were compared with wounding of the bark of 12-15-year-old trees of five conifer species. Wounding in all species resulted in tissue necrosis and wound periderm development immediately around the wound site. One cm from the wound, swelling of phloem polyphenolic parenchyma cells and phenolic accumulation were observed in Pseudotsuga menziesii (Mirb.) Franco, Picea pungens Engelman, Larix occidentalis Nutt. and Pinus monticola Douglas ex D. Don, but not in Taxus brevifolia Nutt. Traumatic resin ducts were formed in response to wounding in three species of Pinaceae, but not in P. monticola, which formed irregular clusters of cells rather than ducts. Taxus brevifolia did not form resin ducts in response to either wounding or MJ treatment. In the Pinaceae species studied, surface application of 100 mM MJ caused similar anatomical changes to those observed in response to wounding, including phenolic accumulation, cell swelling and traumatic resin duct formation, but it did not induce a wound periderm. Traumatic resin ducts differed in size among the study species, ranging from small in L. occidentalis to very large in P. menziesii. In P. menziesii, P. pungens and L. occidentalis, traumatic resin ducts were more abundant after MJ treatment than after wounding. We conclude that the octadecanoid pathway is likely involved in defense responses in stems of the Pinaceae, but not necessarily in other taxa.

Acetates↗

Above- and belowground biomass and net primary production in a 73-year-old Scots pine forest.

We estimated above- and belowground biomass and net primary production (NPP) of a 73-year-old Scots pine (Pinus sylvestris L.) forest stand in the Belgian Campine region. Total biomass for the stand was 176 Mg ha(-1), of which 74.4% was found in stems. The root system contained 12.6% of total biomass, most of it in coarse roots (> 5 mm). Fine roots (< 5 mm) comprised only about 1.7% of total biomass, and more than 50% of fine root biomass was retrieved in the litter layer and the upper 15 cm of the mineral soil. The ratio of belowground biomass to aboveground biomass was 0.14, which is lower than that of other Scots pine forests and other coniferous forests. Between 1995 and 2001, mean annual NPP was 11.2 Mg ha(-1) year(-1), of which 68.7% was allocated to aboveground compartments. Stems, needles and cones made relatively high contributions to total NPP compared with branches. However, branch NPP was possibly underestimated because litterfall of big branches was neglected. The proportion of total NPP in belowground components was 31.3%. Coarse root NPP (2% of total) was low compared with its biomass. Fine root NPP was 3.3 Mg ha(-1) year(-1), representing about 29.5% of total NPP; however, the estimate of fine root NPP is much more uncertain than NPP of aboveground compartments. The ratio NPP/GPP (gross primary production) was 0.32, which was low compared with other coniferous forests.

Belgium↗

Component carbon fluxes and their contribution to ecosystem carbon exchange in a pine forest: an assessment based on eddy covariance measurements and an integrated model.

We used a combination of eddy flux, canopy, soil and environmental measurements with an integrated biophysical model to analyze the seasonality of component carbon (C) fluxes and their contribution to ecosystem C exchange in a 50-year-old Scots pine forest (Pinus sylvestris L.) in eastern Finland (62 degrees 47' N, 30 degrees 58' E) over three climatically contrasting years (2000-2002). Eddy flux measurements showed that the growing Scots pine forest was a sink for CO2, with annual net C uptakes of 131, 210 and 258 g C m-2> year-1 in 2000, 2001 and 2002, respectively. The integrated process model reproduced the annual course of daily C flux above the forest canopy as measured by the eddy covariance method once the site-specific component parameters were estimated. The model explained 72, 66 and 68% of the variation in daily net C flux in 2000, 2001 and 2002, respectively. Modeled annual C loss by respiration was 565, 629 and 640 g C m-2 year-1, accounting for 77, 77 and 65% of annual gross C uptake, respectively. Carbon fluxes from the forest floor were the dominant contributors to forest ecosystem respiration, with the fractions of annual respiration from the forest floor, foliage and wood being 46-62, 27-44 and 9-10%, respectively. The wide range in daily net C uptake during the growing season was largely attributable to day-to-day fluctuations in incident quantum irradiance. During just a few days in early spring and late autumn, ecosystem net C exchange varied between source and sink as a result of large daily changes in temperature. The forest showed a greater reduction in gross C uptake by photosynthesis than in C loss by respiration during the dry summer of 2000, indicating that interannual variability in ecosystem net C uptake at this site was modified mostly by summer rainfall and vapor pressure deficit.

Carbon↗

Growth response and sapwood hydraulic properties of young lodgepole pine following repeated fertilization.

We examined how tree growth and hydraulic properties of branches and boles are influenced by periodic (about 6 years) and annual fertilization in two juvenile lodgepole pine (Pinus contorta Dougl. var. latifolia Engelm.) stands in the interior of British Columbia, Canada. Mean basal area (BA), diameter at breast height (DBH) and height increments and percent earlywood and sapwood hydraulic parameters of branches and boles were measured 7 or 8 years after the initial treatments at Sheridan Creek and Kenneth Creek. At Sheridan Creek, fertilization significantly increased BA and DBH increments, but had no effect on height increment. At Kenneth Creek, fertilization increased BA, but fertilized trees had significantly lower height increments than control trees. Sapwood permeability was greater in lower branches of repeatedly fertilized trees than in those of control trees. Sapwood permeabilities of the lower branches of trees in the control, periodic and annual treatments were 0.24 x 10(-12), 0.35 x 10(-12) and 0.45 x 10(-12) m2 at Kenneth Creek; and 0.41 x 10(-12), 0.54 x 10(-12) and 0.65 x 10(-12) m2 at Sheridan Creek, respectively. Annual fertilization tended to increase leaf specific conductivities and Huber values of the lower branches of trees at both study sites. We conclude that, in trees fertilized annually, the higher flow capacity of lower branches may reduce the availability of water to support annual growth of the leader and upper branches.

Pinus↗

Radial profiles of sap flow with increasing tree size in maritime pine.

We investigated the radial variation of sap flow within sapwood below the live crown in relation to tree size in 10-, 32-, 54- and 91-year-old maritime pine stands (Pinus pinaster Ait.). Radial variations were determined with two thermal dissipation sensors; one measured sap flux in the outer 20 mm of the xylem (Jref), whereas the other was moved radially across the sapwood in 20-mm increments to measure sap flux at multiple depths (Jref). For all tree sizes, sap flow ratios (Ri = JiJref (-1)) declined with increasing sapwood depth, but the decrease was steeper in trees with large diameters. Correction factors (C) were calculated to extrapolate Jref for an estimate of whole-tree sap flux. A negative linear relationship was established between stem diameter and C, the latter ranging from 0.6 to 1.0. We found that neglecting these radial corrections in 10-, 32-, 54- and 91-year-old trees would lead to overestimation of stand transpiration by 4, 14, 26 and 47%, respectively. Therefore, it is necessary to account for the differential radial profiles of sap flow in relation to tree size when comparing tree transpiration and hydraulic properties among trees differing in size.

Pinus↗

Use of modeled photosynthesis and decomposition to describe tree growth at the northern tree line.

Growth of subarctic Scots pine (Pinus sylvestris L.) trees was investigated by a combination of process-based models and dendroecological approaches. Tree ring width indices were strongly autocorrelated and correlated with simulated photosynthetic production of the previous year and with organic matter N mineralization of the current year. An autoregressive model, with photosynthesis and N mineralization as external inputs, explained growth of the trees well. However, relationships for the period 1950-1992 differed significantly from relationships for the period 1876-1949; the slope of the regression of tree ring width index and photosynthesis was lower for the 1950-1992 period. Also, the autocorrelation structure of the data changed. First-order autocorrelation decreased and second-order autocorrelation increased from the earlier to the later period. This means that growth is becoming less sensitive to variations in photosynthetic production, whereas the relationships between growth and N mineralization are remaining fairly constant. We postulate that, although photosynthesis has increased in response to increasing CO2 concentrations, tree growth rate cannot parallel the increase in photosynthesis because potential growth rate is limited directly by temperature.

Carbon Dioxide↗

Acclimation of photosynthetic capacity in Scots pine to the annual cycle of temperature.

Coniferous trees growing in the boreal and temperate zones have a clear annual cycle of photosynthetic activity. A recent study demonstrated that the seasonal variation in photosynthetic capacity of Scots pine (Pinus sylvestris L.) could be attributed mainly to the light response curve of photosynthesis. The magnitude of the light response curve varied over the season while its shape remained constant, indicating that the two physiological parameters quantifying the curve-the quantum yield per unit internal carbon dioxide concentration and the corresponding light-saturated rate-remained proportional to each other. We now show, through modeling studies, that the quantum yield (and hence the light-saturated rate) is related to the annual cycle of temperature through a delayed dynamic response. The proposed model was tested by comparing model results with intensive measurements of photosynthesis and driving variables made from April to October in three shoots of Scots pine growing near the northern timberline. Photosynthetic capacity showed considerable acclimation during the growing season. A single model describing photosynthetic capacity as a reversible, first-order delay process driven by temperature explained most of the variation in photosynthetic capacity during the year. The proposed model is simpler but no less accurate than previous models of the annual cycle of photosynthetic capacity.

Acclimatization↗