Stem growth and respiration in loblolly pine plantations differing in soil resource availability

Stem respiration and growth in 10-year-old loblolly pine (Pinus taeda L.) plantations were measured monthly during the third year of fertilization and irrigation treatments to determine whether soil resource availability differentially altered growth and respiration in stem tissue. Fertilized trees had significantly greater stem biomass, stem nitrogen concentration ([N]) and growth rate than unfertilized trees. Stem respiration (Rt) was significantly greater in fertilized trees when expressed on a per unit surface area (Rt,a, micromol CO2 m-2 s-1), sapwood volume (Rt,v, micromol CO2 m-3 s-1), or mass (Rt,w, nmol CO2 g-1 s-1) basis; however, there was no difference between treatments when expressed as a function of stem N content (Rt,n, micromol CO2 (mol N)-1 s-1). Irrigation had no significant effect on Rt or annual stem growth. Daily total respiration (Rd, mol CO2 m-2 day-1) and stem diameter growth both had a seasonal bimodal pattern with peaks in early spring and midsummer. Stem [N] declined significantly during the growing season. Stem growth rate and [N] explained 75% of the seasonal variation in temperature-normalized Rt,a. The mature tissue method was used to partition total stem respiration (Rt) into maintenance (Rm) and growth (Rg) components. There was a linear correlation between winter Rt,v, a measure of basal Rm, and sapwood N content; however, Rt,v per unit N was greater in January before diameter growth started than in the following December after growth ceased, indicating that Rt,v declined as stem diameter increased. Consequently, estimates of annual maintenance respiration (RM) based on January data were 44% higher than estimates based on December data. Growth respiration was correlated with stem growth rate (r2 = 0.55). The growth respiration coefficient (rg)-the slope of the relationship between Rg and stem growth rate-was 0.24. Respiration accounted for 37% of annual stem carbon budget. Stem carbon-use efficiency (CUE)-the ratio of stem growth to stem growth plus respiration-averaged 0.63 and was unaffected by fertilization.     

Stem form variations in the natural stands of major commercial softwoods in eastern Canada

Stem form is often used as a sawlog assortment criterion and has an important effect on lumber recovery and mechanical properties. Based on 7018 stems collected from the natural stands in 3 regions in eastern Canada, this study quantified the variations in stem form (taper, sweep and eccentricity) for five major commercial softwood species, viz., jack pine (Pinus banksiana Lamb.), black spruce (Picea mariana), white spruce (Picea glauca), red spruce (Picea rubens) and balsam fir (Abies balsamea). The majority of the stems had a diameter at breast height (DBH) ranging from 10 to 32 cm and total tree height from 10 to 20 m. Stem taper and butt taper increased steadily from 0.38 to 1.75 cm/m and from 0.74 to 4.23 cm/m, respectively, with DBH increasing from 8 to 48 cm. Stem sweep and basal sweep ranged from 0.12 to 7.28 cm/m and from 0.12 to 9.85 cm/m, respectively, with mean values of 0.87 and 0.95 cm/m. There were no significant differences in stem sweep and basal sweep between DBH classes and both sweeps distributed over a wide range within DBH classes, species and regions. About 0.77% of the stems and 1.2% of the butt logs had seriously curved shapes (sweep >3.0 cm/m). Eccentricity (ratio of smaller diameter to larger diameter) at the breast height (0.96) was very close to 1. Balsam fir and jack pine had the best and worst stem forms, respectively, while stem forms in black spruce were in-between. Butt logs were most seriously curved in red spruce and most tapered in white spruce. Region C (mainly eastern Quebec and the Atlantic Provinces) produced stems with smaller butt taper, basal sweep and stem sweep and larger eccentricity compared with other regions in eastern Canada. Overall stems in eastern Canada have a good stem form in terms of taper, sweep and eccentricity. The evaluation of stem form provides valuable timber quality information for the wood industry to optimize wood processing as well as for forest managers to define appropriate silvicultural regimes to improve stem form and economic value.     

Stem deformation in young plantations of black spruce (Picea mariana (Mill.) B.S.P.) and jack pine (Pinus banksiana Lamb.) in the boreal forest of Quebec,Canada

Stem deformation has often been observed in young black spruce (Picea mariana (Mill.) B.S.P.) and jack pine (Pinus banksiana Lamb.) plantations. Whenever important stem deformations are observed at the time of harvesting, timber value is negatively affected especially during the wood transformation process. The present work was undertaken to quantify and qualify the importance of stem deformation of black spruce and jack pine in the boreal forest of central Quebec at the stand and tree levels. In 30 black spruce and jack pine plantations, approximately 22% of spruce trees and 27% of pine trees exhibited stem deformation. The proportion of deformed trees was higher in the youngest plantations and decreased with the age of the plantations. Stem deformation caused the formation of compression wood which is another factor that can reduce the value of wood products. Thirty-nine black spruces and 34 jack pines were analysed at the tree level. On average, compression wood represented 14% and 20% of stem volume in 7- and 10-year old black spruce plantations, respectively. These proportions ranged from 18% in the youngest jack pine plantation to 26% in the oldest one. Stems of both species classified as normal contained a lower volume of compression wood than stems classified as deformed or very deformed. Annual percentages of compression wood and annual shoot length increased significantly with tree age (p < 0.0001 for both variables). Statistically significant correlations were also found between the range of displacement of the stem and the percentage of compression wood. The fewer number of trees with deformed stems in older plantations combined with high compression wood formation suggests that, over time, a deformed tree can become normal and straight in appearance.     

Relation between wilt rate and obstruction of water flow in stems of two families of loblolly pine affected by pitch canker

Movement of water forced through stem segments of two half-sib families of loblolly pine (Pinus taeda), infected by Fusarinum subglutinans, was monitored. Wilt rate of the two families was shown to be correlated with obstruction of water flow through stem segments of similar disease severity.     

Respiratory responses of Scots pine stems to 5 years of exposure to elevated CO2 concentration and temperature

Stem respiration in 20-year-old Scots pine (Pinus sylvestris L.) trees was examined following 5 years of exposure to ambient conditions (CON), elevated atmospheric carbon dioxide concentration ([CO2]) (ambient + 350 micromol mol(-1), (EC)), elevated temperature (ambient + 2-6 degrees C, (ET)) or a combination of elevated [CO2] and elevated temperature (ECT). Stem respiration varied seasonally regardless of the treatment and displayed a similar trend to temperature, with maximum rates occurring around Day 190 in summer and minimum rates in winter. Respiration normalized to 15 degrees C (R15) was higher in the growing season than in the non-growing season, whereas the temperature coefficient (Q10) was lower in the growing season. Annually averaged R15 was 0.36, 0.43, 0.40 and 0.44 micromol m(-2) s(-1) under CON, EC, ET and ECT conditions, respectively, whereas the corresponding values for total stem respiration were 6.55, 7.69, 7.50 and 7.90 mol m(-2) year(-1). The EC, ET and ECT treatments increased R15 by 18, 11 and 22%, respectively, relative to CON, and increased the modeled annual total stem respiration by 18, 15 and 21%. The increase in modeled annual stem respiration under EC and ECT conditions was caused mainly by higher maintenance respiration (22 and 25%, respectively, whereas the increase in growth respiration was 9 and 12%). Growth respiration was unaltered by ET. The treatments did not significantly affect the respiratory response to stem temperature; the mean Q10 value was 2.04, 2.10, 1.99 and 2.12 in the CON, EC, ET and ECT treatments, respectively. It is suggested that the increase in stem respiration was partly a result of the increased growth rate. We conclude that elevated [CO2] increased the maintenance component of respiration more than the growth component.     

Irradiance in thinned Norway spruce (Picea abies) stands and the possibilities to prevent suckers of broadleaved trees

Both incoming shortwave radiation (R_g) and photosynthetically active radiation (PAR) in percentage of full daylight were measured at the same time by point and strip sampling in four plots (0.1 ha) of Picea abies (L.) Karst. The standard deviations (%) of R_g and PAR were, respectively, 11.1 and 9.8 at 64 points, 15.7 and 13.9 at 32 points, and 24.7 and 23.8 at 16 points per plot.

A period of at least 40 s per strip (30 m min⁻¹) gives a CV (coefficient of variation) of 30%. There is no significant difference between relative irradiance (RI) estimated by the point method (64 points) and by the strip method (8 strips). Curves of RI (R_g and PAR) and basal area (m² ha⁻¹), diameter sum (m ha⁻¹) and density (stems ha⁻¹) of fifteen trials with different thinning programmes are presented. Irradiance (R_g) in heavily thinned stands was 3–14% of irradiance on an open place. The irradiance, R_g, in extra-heavily thinned stands is 12–27%, and in unthinned stands, 1–3% that of an open place. The R_g curve lies above the PAR curve in all cases. Some practical implications of the study are presented. Heavy thinning of Norway spruce stands gives RI (R_g) values 10% at basal area of 25m² ha⁻¹ which is necessary to minimize development of suckers of broadleaved trees.     

Stem respiration is influenced by pruning and girdling in Pinus sylvestris

Abstract

Stem respiration was measured in the growing season (June to July) and in the dormant season (October) to detect cambial activity induced by pruning live branches or girdling stems in Scots pine trees (Pinus sylvestris L.) growing in northern Sweden. Immediately after the treatments, the treatment:control ratio of stem respiration increased to between 1.38 and 1.44 in the pruning treatment and between 1.17 and 1.20 in the girdling treatment. The treatment:control ratio of stem respiration then decreased by the end of July, to 0.65 in the pruning treatment and 0.55 in the girdling treatment. In October, the treatment:control ratios were higher: between 0.87 and 0.97 in the pruning treatment and between 0.85 and 0.97 in the girdling treatment. In both pruning and girdling treatments, the time trends of stem respiration rates largely followed those of stem temperatures: the stem respiration rate increased exponentially with an increase in stem temperature. The Q ₁₀ values were 2.83–4.05 and 2.57–2.89 in the pruning treatment and control, and 2.10–2.60 and 1.99–3.19 in the girdling treatment and control, respectively. In most cases, the values of Q ₁₀ in both treatments did not differ significantly from those in the controls.     

Effects of waterlogging on the growth and water relations of three Pinus taxa

The response of Caribbean pine (Pinus caribaea Morelet. var. hondurensis Barr. and Golf.), slash pine (Pinus elliottii Englem. var. elliottii) and their F₁ hybrid to soil waterlogging was studied under glasshouse conditions. Plants were grown in either a lateritic or gleyed podzolic soil and pots were flooded in both autumn and summer treatments.

Plant responses were influenced by both season and duration of flooding, and by soil type. All the Caribbean pine seedlings survived the autumn flooding, but showed very slow growth and had very low needle conductances. Summer-flooded plants exhibited rapid stomatal closure and low needle water-potentials accompanied by needle dehydration. All of the summer-flooded Caribbean pine seedlings died. Slash pine was much more flood-tolerant than was Caribbean pine, whilst the F₁ hybrid was intermediate between the two parents. All slash pine and F₁ hybrid seedlings survived autumn flooding and continued to grow, but survival was reduced with summer flooding, particularly in the short-term flooding treatment. Slash pine and F₁ hybrid seedlings showed morphological features of flood tolerance with production of adventitious roots and formation of lenticels in association with stem hypertrophy. Although stomatal opening was evident during autumn flooding, needle conductance was very much reduced in summer-flooded plants of these two taxa. Needle water-potential and relative water content were maintained near the control value in flooded slash pine grown in both soil types, but only for the F₁ hybrid in the flooded lateritic podzolic soil.     

Comparison of field methods for measuring soil respiration: a static alkali absorption method and two dynamic closed chamber methods

The objectives of the present study were to compare the static alkali absorption (AA) and dynamic closed chamber (DC) methods for measuring soil respiration, and to evaluate the effects of methodological differences on estimating annual mean soil respiration rate in a natural forest. For the AA method, we used Kirita’s method [Jpn. J. Ecol. 21 (1971) 119] using an alkali-soaked sponge disc that covers nearly the same area as that covered by a chamber. For the DC method, we used both the LI-6200 system (DC-62 method) and the newer LI-6400 system (DC-64 method) (LI-COR, Lincoln, NE, USA). Comparative measurements were conducted on five occasions during the study period (November 1998–October 1999) at a Quercus serrata forest in Japan. Daily mean soil respiration rates obtained by the AA, DC-62 and DC-64 methods for a 24 h period were in the ranges 205–578, 147–629 and 165–734 mg CO₂ m⁻² h⁻¹, respectively. The daily mean soil respiration rates obtained by the AA method were 79–128% of those obtained by the DC-64 method. When the daily mean soil respiration rate obtained by the DC-64 method was below 300 mg CO₂ m⁻² h⁻¹, the daily mean soil respiration rate obtained by the AA method was an average of 26% higher than that obtained by the DC-64 method. When the daily mean soil respiration rate obtained by the DC-64 method was above 300 mg CO₂ m⁻² h⁻¹, the daily mean soil respiration rate obtained by the AA method was an average of 19% lower than that obtained by the DC-64 method. However, at the present site, there was a little difference between the two methods as for estimating annual mean soil respiration rate, and therefore the AA method improved by Kirita [Jpn. J. Ecol. 21 (1971) 119] is suggested to be a useful method for estimating annual mean soil respiration in the forest. The daily mean soil respiration rates obtained by the DC-62 method were systematically 10–24% lower (an average of 15% lower) than those obtained with the DC-64 method, and the annual mean rate was lower than that estimated by the AA method.     

Comparison of organic matter dynamics in soil between Japanese cedar (Cryptomeria japonica) forest and adjacent Japanese red pine (Pinus densiflora) forest established on flatland

In order to clarify the effects of tree species on organic matter dynamics in soil, we investigated the amount of forest floor material, leaf litter decomposition rate, soil chemical characteristics, soil respiration rate and cellulose decomposition rate in a Japanese cedar forest (cedar plot) and an adjacent Japanese red pine forest (pine plot) established on a flatland. The amount of forest floor material in the cedar plot was 34.5 Mg ha⁻¹ which was greater than that in the pine plot. Because the leaf litter decomposition rate was higher in the pine plot than in the cedar plot, it is likely that the difference in the amount of forest floor material between the plots is caused by the difference in the leaf litter decomposition rate. The C concentrations of soil in the cedar plot were 1.2–2.1 times higher than those in the pine plot. Soil pH(H₂O)s in the cedar plot were significantly higher than those in the pine plot. The soil respiration rates and the rates of mineralized C in the cedar plot byin vitro incubation were higher than those in the pine plot. From this result, it is assumed that soil organic matter in the cedar plot was decomposed relatively faster compared with the pine plot. Furthermore, microbial activities, which were reflected as cellulose decomposition rates in the cedar plot, were higher than those in the pine plot. A part of this paper was presented at the 109th Annual Meeting of the Japanese Forestry Society (1998).     

Growth responses, physiology and decay associated with pruning plantation-grown Eucalyptus globulus Labill. and E. nitens (Deane and Maiden) Maiden

Detailed studies have been undertaken to define pruning regimes for Eucalyptus nitens, but little is known of E. globulus responses to pruning although this is a more commonly planted species. This paper describes experiments that aimed to identify (a) levels of pruning that reduce E. globulus growth, (b) physiological processes contributing to those growth responses, (c) the incidence of decay and factors influencing decay incidence following pruning of E. globulus, and (d) comparative responses of E. nitens and E. globulus to live branch pruning. Results of a field experiment indicated that removal of between 30 and 50% of the crown length was appropriate for E. globulus plantations verging on canopy closure. The significant reduction in height growth associated with removal of 50 or 70% of crown length suggested pruning should remain below 50% of crown length if reduced stem growth of pruned trees was to be avoided. Stem volume was only significantly reduced over the period of the experiment by 70% pruning, but it was estimated that standing volume following removal of 50% of crown length would be reduced by 82 m³ ha⁻¹ over a 20-year rotation if there were no other silvicultural interventions. The growth responses observed were probably related to large reductions in leaf area following 50 or 70% removal of crown length. Trees responded to pruning by changing patterns of biomass partitioning to favour stem growth at the expense of branch growth. A glasshouse study determined that light-saturated net CO₂ uptake (A_max) increased following pruning. E. nitens seedlings had both a higher baseline A_max and higher A_max following pruning than did E. globulus, which could partially explain the greater effect of pruning on E. globulus growth than has been observed for E. nitens in other studies. This result, as well as apparently different patterns of foliage distribution through the crowns of E. globulus and E. nitens, suggested that models of pruning responses need to be parameterised for both species. In addition, a more conservative pruning regime may be appropriate for E. globulus than E. nitens. Pruning increased the frequency of branch traces with decay infection, and there was a trend towards increasing decay outbreaks with increasing pruning severity. Decay outbreaks were more likely to occur following pruning of high angle or larger diameter branches.     

The effect of aqueous transport of CO(2) in xylem sap on gas exchange in woody plants

The influence of CO(2) transported in the transpiration stream on measurements of leaf photosynthesis and stem respiration was investigated. Measurements were made on trees in a temperate forest in Scotland and in a tropical rain forest in Cameroon, and on shrubs in the Sahelian zone in Niger. A chamber was designed to measure the CO(2) partial pressure in the gas phase within the woody stems of trees. High CO(2) partial pressures were found, ranging from 3000 to 9200 Pa. Henry's Law was used to estimate the CO(2) concentration of xylem sap, assuming that it was in equilibrium with the measured gas phase partial pressures. The transport of CO(2) in the xylem sap was calculated by multiplying sap CO(2) concentration by transpiration rate. The magnitude of aqueous transport in the studied species ranged from 0.03 to 0.35 &mgr;mol CO(2) m(-2) s(-1), representing 0.5 to 7.1% of typical leaf photosynthetic rates. These values strongly depend on sap pH. To examine the influence of aqueous transport of CO(2) on stem gas exchange, we made simultaneous measurements of stem CO(2) efflux and sap flow on the same stem. After removing the effect of temperature, stem CO(2) efflux was positively related to sap flow. The apparent effect on measurements of stem respiration was up to 0.7 &mgr;mol m(-2) s(-1), representing ~12% of peak stem respiration rates.     

Stem Form and Taper Changes After Thinning and Nitrogen Fertilization in Picea abies and Pinus sylvestris Stands

Stem form and taper changes after thinning and thinning combined with N fertilization were studied in 23 Norway spruce [Picea abies (L.) Karst.] and 46 Scots pine (Pinus sylvestris L.) stands in Sweden observed for 5-28 yrs. Average taper was calculated for the stem section 1.3-6.0 m above ground level and as the ratio between the diameter at breast height and total tree height for periods of 5-10 yrs. At the start of the experiment, before the first thinning, the dominant height was 12-15 m. Once 65% of the basal area had been removed in a single thinning from below, the remaining trees showed a strong increase in taper compared with trees in unthinned stands. Stems in stands treated with heavily recurrent thinnings from below also increased in taper, although the increases were not always statistically significant. Trees in thinned, N-fertilized Scots pine stands in middle and northern Sweden developed a more pronounced taper compared with stems in equally thinned, unfertilized stands.     

Estimation of whole-stem respiration,incorporating vertical and seasonal variations in stem CO<Subscript>2</Subscript> efflux rate,of <Emphasis Type="Italic">Chamaecyparis obtusa</Emphasis> trees

We examined vertical and seasonal variations in stem respiration rates in a 50-year-old plantation of Japanese cypress, Chamaecyparis obtusa (Sieb. et Zucc.) Endl., in central Japan, and discuss a practical and precise method to scale a point-measured stem CO₂ efflux rate up to whole-stem respiration. For five selected trees, stem CO₂ efflux rates were measured at breast height (1.3 m) and at five or six points above breast height (at approximately 2 m intervals) every 1 or 2 months over two consecutive years. Daily total stem respiration rate (surface area basis) was greater inside the crown than below the crown, especially during the growing season. By incorporating the vertical profile of the respiration rate, annual whole-stem respiration was estimated for each sample tree (R _y). We then compared this estimate (R _y) with another estimate of annual whole-stem respiration (R′_y) obtained using a conventional method; it is assumed that the area-based respiration rate at breast height is constant throughout the stem. The ratio of these two estimates (R′_y/R _y) was usually less than 1, indicating that the assumptions used to calculate R′_y underestimate annual whole-stem respiration. We found that R′_y/R _y was negatively correlated with the ratio of crown length to tree height (crown ratio). These results suggest that annual whole-stem respiration in this C. obtusa plantation is substantially affected by the relative proportion of within-crown stem with higher respiratory activity. Methodologically, our results imply that incorporating the crown ratio into the conventional method would improve the accuracy of annual whole-stem respiration estimates.     

Effects of the interaction between drought and shade on water relations, gas exchange and morphological traits in cork oak (Quercus suber L.) seedlings

The combined effect of drought and light on different physiological and biochemical traits was assessed in cork oak (Quercus suber L.) seedlings grown under two levels of light availability and submitted to a long-standing drought. Watering was withdrawn after germination and seedlings were allowed to dry to a water content of ca. 50% of field capacity. At this point, water-stressed seedlings were grown under moderate drought and two light regimes: high light (HL—50%) and low light (LL—2%). Soil water in control plants was kept close to field capacity (90–100%) for both light environments. Water-relations parameters derived from P–V curves, gas exchange and water status at predawn (Ψ_pd) were evaluated at twice during the experiment. Nitrogen and chlorophyll contents were determined in the same leaves used for the gas exchange measurements. In addition, maximum rate of carboxylation (V_cmax) and electronic transport (J_max) were derived from A–C_i curves in well-watered seedlings.

The variation on moisture availability during the experiment was the same under both light environments. In control plants, Ψ_pd was over −0.3 MPa at the two harvests, while stressed seedlings decreased to −0.9 MPa, with no differences between light treatments. Water stress decreased osmotic potentials at full (Ψπ₁₀₀) and zero turgor (Ψπ₀). The regressions between both potentials and Ψ_pd showed a higher intercept in shade grown seedlings. This fact will point out the higher osmoregulation capacity in sun seedlings whatever water availability.

Nitrogen investment on a per leaf mass (N_mass), chlorophyll content (Chl_mass) and SLA tended to show a typical pattern of sun-shade acclimation. Thus, the three parameters increased with shade. Only for N_mass there was a significant effect of watering, since water stress increased N_mass.

LL plants showed a lower photosynthetic capacity in terms of maximum net photosynthesis at saturating light (A_max), which was related to a decrease in V_cmax and J_max. Both parameters varied with specific leaf area (SLA) in a similar way. The low-light environment brought about a higher nitrogen investment in chlorophyll, while under high-light environment the investment was higher in carboxylation (V_cmax) and electronic transport (F_max).

Stomatal conductance to water vapour (g_wv) and A_max were lower in low-light seedlings independently of watering. In addition, there was a trend to keep higher intrinsic water use efficiency (IWUE) under high light environment. The increase of IWUE under water stress was higher in HL seedlings. This was as consequence of the steeper decline in g_wv as Ψ_pd decreased. The decrease of A_max with Ψ_pd occurred in a similar way in LL and HL seedlings. Thus, the HL seedlings tended to sustain a higher ability to increase IWUE than LL seedlings when they were submitted to the same water stress.     

Influence of plantation site and wastewater sludge fertilization on the performance and foliar nutrient status of two willow species grown under SRIC in southern Quebec (Canada)

Salix discolor Mühl. (Sd) and Salix viminalis L. (Sv) were planted under short-rotation intensive culture (SRIC) on three unirrigated abandoned farmland sites with different drainage conditions, one well-drained (S₁) and the other two poorly-drained (S₂, S₃). One dose of dried and granulated sludge equivalent to 150 (T₁) kg of “available” N ha⁻¹ was applied to some plots in the spring of the second season while others were left unfertilized (T₀). The aims of the experiment were (i) to investigate plant response (growth and productivity) to plantation site conditions and sludge application; (ii) study nutrient status by foliar analysis. Over three seasons, growth in height and aboveground biomass were greater for S. viminalis than for S. discolor on all sites. S. viminalis planted on poorly-drained site 2 had the highest biomass yield (45.28 t ha⁻¹). Both species showed best height and diameter growth on poorly-drained sites. For both species, best performances were obtained on wastewater sludge fertilized plots. Comparative foliar analysis suggested that unfertilized sandy soil (S₁) and low foliar nitrogen concentration and content were limiting factors in the performance of the two species. Soil nitrate concentration increased as a result of sludge application. Heavy metal accumulation from sludge does not represent a risk to the environment. It was concluded that S. viminalis had the best productivity on clay sites, and that a moderate dose of dried and pelleted sludge (150 kg of “available” N ha⁻¹) may be a good fertilizer during the establishment of willows in SRIC, and may reduce nitrate leaching.     

Nitrogen mineralization along an altitudinal gradient: Interactions of soil temperature, moisture, and substrate quality

Net soil N mineralization was measured in situ for 2 consecutive years along a 2000-m altitudinal gradient encompassing six vegetation types in northern California. Both anaerobic and aerobic field incubations showed that soil temperature and moisture strongly controlled N release. In sealed container studies, N mineralization per unit of total Kjeldahl N ( ) had a Q₁₀ rate of 2 and varied between 5 and 38 g N/kg when moisture was abundant (anaerobic incubation). However, aerobic mineralization rates fell to between 2 and 22 g N/kg because of summer drought. Aerobic rates were greatest at mid-elevations (the mixed-conifer forest), and were reduced by cold temperatures at higher elevations and by soil drought at lower. The lowest rate found for any site was in a pine plantation scalped of topsoil a decade previously, during site preparation. Open-container incubations involving intact, 15-cm soil cores and ion-exchange resins produced mineralization rates correlated with potential growing days, but not with rates from sealed containers. The forest floor accounted for nearly all the net N mobilized on the coldest site, but only one-half to one-third of the N mobilized on warmer sites. Open containers have important advantages in assessing soil N dynamics.     

Effects of nitrogen on the growth of jack pine competing with Canada blue-joint grass and large-leaved aster

Biomass, leaf area, canopy photosynthesis, photosynthetic nitrogen-use efficiency (PNUE), nitrogen-partitioning ratio (NPR: ratio of nitrogen taken up by jack pine relative to two different competitor species), and nitrogen uptake (NU) of jack pine (Pinus banksiana Lamb.) competing with large-leaved aster (Aster macrophyllus L.) and Canada blue-joint grass (Calamagrostis canadensis (Michx.) Beauv.) were examined at three nitrogen levels in a controlled-environment growth chamber. When grown with large-leaved aster, jack pine biomass, photosynthesis and PNUE (p<0.001) increased as nitrogen level increased. Jack pine biomass, photosynthesis and NPR (p<0.001) decreased as nitrogen level increased when grown with Canada blue-joint grass. At the lowest nitrogen supply level, jack pine photosynthesis decreased as competitor PNUE increased (r²=0.84, p<0.001). Jack pine photosynthesis decreased as NU of large-leaved aster (37.5 mg N l⁻¹: r²=0.75, p<0.001; 100 mg N l⁻¹: r²=0.86, p<0.001) and Canada blue-joint grass (37.5 mg N l⁻¹: r²=0.96, p<0.001; 100 mg N l⁻¹: r²=0.84, p<0.001) increased. NU and PNUE may play an important role in the outcome of interactions between jack pine seedlings and competing forest vegetation in newly planted stands.     

Predicting the decline in daily maximum transpiration rate of two pine stands during drought based on constant minimum leaf water potential and plant hydraulic conductance

The effect of drought on forest water use is often estimated with models, but comprehensive models require many parameters, and simple models may not be sufficiently flexible. Many tree species, Pinus species in particular, have been shown to maintain a constant minimum leaf water potential above the critical threshold for xylem embolism during drought. In such cases, prediction of the relative decline in daily maximum transpiration rate with decreasing soil water content is relatively straightforward. We constructed a soil-plant water flow model assuming constant plant conductance and daily minimum leaf water potential, but variable conductance from soil to root. We tested this model against independent data from two sites: automatic shoot chamber data and sap flow measurements from a boreal Scots pine (Pinus sylvestris L.) stand; and sap flow measurements from a maritime pine (Pinus pinaster Ait.) stand. To focus on soil limitations to water uptake, we expressed daily maximum transpiration rate relative to the rate that would be obtained in wet soil with similar environmental variables. The comparison was successful, although the maritime pine stand showed carry-over effects of the drought that we could not explain. For the boreal Scots pine stand, daily maximum transpiration was best predicted by water content of soil deeper than 5 cm. A sensitivity analysis revealed that model predictions were relatively insensitive to the minimum leaf water potential, which can be accounted for by the importance of soil resistance of drying soil. We conclude that a model with constant plant conductance and minimum leaf water potential can accurately predict the decline in daily maximum transpiration rate during drought for these two pine stands, and that including further detail about plant compartments would add little predictive power, except in predicting recovery from severe drought.