Growth and physiological response of eastern white pine seedlings to partial cutting and site preparation

We examined how white pine (Pinus strobus L.) seedlings planted under a mature cover of white and red (Pinus resinosa Ait.) pine in eastern Ontario (Canada) responded to treatments aimed at improving light and soil conditions for seedling growth. The treatments were: (a) three levels of partial cutting (no cut or CS0, cut to one-crown spacing between residual trees or CS1, cut to two-crown spacing or CS2); (b) two levels of vegetation control (without herbicide or H0, with herbicide or H1); and (c) two levels of soil scarification (S0 and S1). On the third growing season after planting, total growth of seedlings was lowest in CS0 treatment and similar in CS1 and CS2 treatments. The CS2 created better growing light conditions than the CS1, with and average of 50% of full light at seedling height, which corresponded to the maximum height and diameter growth rates of seedlings. However, CS2 also stimulated the growth of competing woody vegetation (both understory trees and shrubs), and resulted in greater microsite heterogeneity of light availability. Scarification warmed the soil (approximately 1–3 °C in the middle of the growing season), decreased the density of competing trees, but increased the shrub density, with no impact on white pine seedling growth. The treatments had no effect on light-saturated photosynthetic rate (A) of current-year foliage of seedlings, nor on their midday shoot water potential. Leaf N was higher in partial cuts and with vegetation control, but the relationship between N and A was weak to non-existent for the different foliage classes. Measures of the proportion of aboveground biomass allocated to foliage (leaf-mass ratio) suggest an acclimation response of young white pine that improves growth under moderate light availability and compensates for the lack of leaf-level photosynthetic plasticity. We suggest a combination of soil scarification under a one-crown spacing partial cut (corresponding to 14 m² ha⁻¹ of residual basal area, or an average of 32% of available light at seedling height) as an establishment cut. This should provide optimum growth conditions for planted understory white pine, while also favoring natural regeneration and providing some protection against damage from insects and disease.     

Effects of soil temperature on biomass production and allocation in seedlings of four boreal tree species

One-year old seedlings of trembling aspen (Populus tremuloides Michx.), black spruce (Picea mariana (Mill.) B.S.P.), white spruce (Picea glauca (Moench) Voss), and jack pine (Pinus banksiana Lamb.) were subject to seven soil temperatures (5, 10, 15, 20, 25, 30 and 35 °C) for 4 months. All aspen seedlings, about 40% of jack pine, 20% of white spruce and black spruce survived the 35 °C treatment. The seedlings were harvested at the end of the fourth month to determine biomass and biomass allocation. It was found that soil temperature, species and interactions between soil temperature and species significantly affected root biomass, foliage biomass, stem biomass and total mass of the seedling. The relationship between biomass and soil temperature was modeled using third-order polynomials. The model showed that the optimum soil temperature for total biomass was 22.4, 19.4, 16.0 and 13.7 °C, respectively, for jack pine, aspen, black spruce and white spruce. The optimum soil temperature was higher for leaf than for root in jack pine, aspen and black spruce, but the trend was the opposite for white spruce. Among the species, aspen was the most sensitive to soil temperature: the maximum total biomass for aspen was about 7 times of the minimum value while the corresponding values were only 2.2, 2.4 and 2.3 times, respectively, for black spruce, jack pine and white spruce. Soil temperature did not significantly affect the shoot/root (S/R) ratio, root mass ratio (RMR), leaf mass ratio (LMR), or stem mass ratio (SMR) (P>0.05) with the exception of black spruce which had much higher S/R ratios at low (5 °C) and high (30 °C) soil temperatures. There were significant differences between species in all the above ratios (P<0.05). Aspen and white spruce had the smallest S/R ratio but highest RMR while black spruce had the highest S/R but lowest RMR. Jack pine had the highest LMR but lowest SMR while aspen had the smallest LMR but highest SMR. Both LMR and SMR were significantly higher for black spruce than for white spruce.     

Density-dependent effects of northern hardwood competition on selected environmental resources and young white spruce (Picea glauca) plantation growth, mineral nutrition, and stand structural development – a 5-year study

The effects of competition from red raspberry (Rubus idaeus L.) and northern hardwood tree species on white spruce (Picea glauca (Moench) Voss) seedlings were examined on a clearcut site of the boreal mixedwood forest of the Bas-Saint-Laurent region of Quebec, Canada. A controlled experiment involving a gradient of five vegetation densities on the basis of the leaf area index (LAI) was established in a completely randomized plot design with six replications. Each of the five levels of vegetation cover (including vegetation-free plots) were examined to evaluate how they affected environmental factors (quantity and quality of light reaching the spruce seedlings, and soil temperature), spruce growth (height, basal diameter, volume index, and above-ground biomass), spruce mortality, browsing damage, spruce foliar mineral nutrition, as well as the stand structural development, during the first 5 years after seedling planting.

Each spruce growth variable analyzed in this study, according to a RMANOVA procedure, followed a negative hyperbolic form of density dependence of competitive effects. Loss of growth in young white spruce plantations in competition with northern hardwoods is likely to occur with the first few competitors. In cases where higher levels of competing vegetation were maintained over time, loss of spruce growth was extremely severe, to an extent where the exponential growth character of the young trees has been lost. At the end of the fifth year, spruce growing with no interference were larger in mean total above-ground biomass by a factor of 9.7 than those growing with the highest level of vegetation cover. Spruce did not develop a strategy of shade avoidance by increasing tree height, on the contrary. Spruce mortality differed among treatments only in the fifth year, indicating that early evaluation of spruce survival is not a strong indicator of competitive effects, when compared to diameter growth. Spruce foliar N and Ca contents were significantly reduced by the first level of competing vegetation cover, while K increased with the density of the vegetation cover, and P and Mg were not affected. Nitrogen nutrition of young white spruce planted on recently disturbed sites is discussed in relation to the potential root discrimination of this species against soil nitrate, a reaction observed by Kronzucker et al. [Kronzucker, H.J., Siddiqi, M.Y., Glass, A.D.M., 1997. Conifer root discrimination against soil nitrate and the ecology of forest succession. Nature London 385, 59–61]. The effects of hardwood competition indicate a prevalence of competition for light over a competition for nutrients, as revealed by the substantial increase in the h/d ratio of white spruce. Two indicators, h/d ratio and the quantity of light received at the tree seedling level, are suggested as a basis for the management of hardwood competition in a white spruce plantation.

Analysis of the stand structural development indicates that spruce height distribution was affected only by moderate or dense cover of vegetation, while diameter distribution, when compared to competing vegetation-free plots, was affected by the lowest level of vegetation cover. This study shows that competition influenced the stand structural development in the same way as genetic and micro-site factors by aggravating the amplitude of size inequality. The impact of hardwood competition is discussed in view of reaching an equilibrium between optimal spruce plantation growth and benefits from further silvicultural treatments, and maintaining hardwood species known to improve long term site quality, within a white spruce plantation.     

Effects of feathermoss removal, thinning and fertilization on lodgepole pine growth, soil microclimate and stand nitrogen dynamics

The influence of a continuous feathermoss layer (dominated by Pleurozium schreberi) on soil temperature, soil water potential and tree growth was assessed in a 5-year study. The feathermoss layer was manually removed from 900 m² plots in lodgepole pine (Pinus contorta) stands of northwestern Alberta. The interception and retention of nutrients by moss was estimated in N fertilization treatments with and without moss removal. As well, the potential for intraspecific competition to affect nutrient dynamics was assessed through a thinning treatment. Removal of the moss layer increased soil temperatures in summer and increased the period when soil was frost free, but the plots without moss had soil temperature as low as −13 °C in one winter period with little snow cover. Moss removal did not significantly affect N concentrations of the tree foliage but did reduce needle weight. Thinning had little effect on the rate of diameter growth after the first 3 years, but produced a significant increase in growth by year 4. Fertilization had a consistently positive effect on radial stem increments and N content of foliage, regardless of the presence of moss on the plot. Overall, the intraspecific competition between trees was apparently greater than interspecific competition between moss and trees.     

Facilitative and competitive effects of a N-fixing shrub on white fir saplings

In Sierra Nevada forests, shrubs are considered strong soil moisture competitors with regenerating trees, reducing seedling establishment, and slowing growth. Recent studies, however, suggest that in some circumstances shrubs can facilitate tree establishment and growth by modifying harsh microclimate conditions; increasing acquisition of water, carbon, and/or nutrients via shared mycorrhizal connections; or enhancing soil fertility, particularly under nitrogen-fixing shrubs such as Ceanothus spp. We examined the establishment dates and growth rates and patterns of white fir saplings growing in greenleaf manzanita, whitethorn ceanothus, and bare patches to examine whether establishment was correlated with past wet years, whether saplings growing in ceanothus had nitrogen-enriched foliage or faster growth rates than in the other two patches, and whether saplings in shrub patches experienced competition for light.Sapling establishment was not correlated with high precipitation or heavy snowpack years, suggesting shade-tolerant saplings do not need wet years to become established. Soils under ceanothus were nitrogen enriched, but white fir sapling foliage did not have higher nitrogen concentrations and saplings did not grow faster in ceanothus than in the other two patches. Because growth rates of saplings were comparable in all patch types examined despite significantly different edaphic and abiotic conditions, we inferred that the various competitive and facilitative interactions affecting tree growth are in net balance across the patch types examined. However, competition for light is important—a significant percentage of growth release events occurred after saplings emerged above their host shrubs. Where shrubs are present, shade-tolerant species (i.e., white fir) are favored over drought-tolerant (pine) species. Our results may help interpret changes in understory conditions that are contributing to mixed conifer's compositional shift toward more shade-tolerant species after a century of fire-suppression.     

Seasonal nutrient dynamics in white pine and white spruce in response to environmental manipulation

Seasonal retranslocation in white pine (Pinus strobus L.) and white spruce (Picea glauca (Moench) Voss) was examined in response to silvicultural treatments (scarification, annual fertilization application, and annual control of competing vegetation with herbicide) that changed both environmental conditions and the growth rate of the trees. Four years after plantation establishment and initial treatment, nutrient accumulation in current-year needles of white pine and retranslocation from 1-year-old needles were increased following the vegetation control treatment, which increased resource availability (nutrients, water and light) and, hence, growth rate. Nutrient accumulation also increased in current-year white spruce needles following the same treatment, whereas retranslocation decreased in 1-year-old white spruce needles. Correlations of retranslocation (N, P and K) with growth rate (shoot biomass increment) showed a strong positive relationship for white pine and a negative relationship for white spruce. Retranslocation of K was correlated with foliar and soil K concentrations; the availability of this nutrient was also significantly reduced by vegetation control. A general theory for the control of nutrient retranslocation in conifers, which is not based exclusively on either sink strength or soil nutrient availability, is proposed. We conclude that retranslocation response is species specific and related to the potential phenotypic growth response to changing environmental conditions and to short-term imbalance in the supply versus the demand for nutrients.     

Interactive effects of elevated ozone and temperature on carbon allocation of silver birch (Betula pendula) genotypes in an open-air field exposure

In the present experiment, the single and combined effects of elevated temperature and ozone (O(3)) on four silver birch genotypes (gt12, gt14, gt15 and gt25) were studied in an open-air field exposure design. Above- and below-ground biomass accumulation, stem growth and soil respiration were measured in 2008. In addition, a (13)C-labelling experiment was conducted with gt15 trees. After the second exposure season, elevated temperature increased silver birch above- and below-ground growth and soil respiration rates. However, some of these variables showed that the temperature effect was modified by tree genotype and prevailing O(3) level. For instance, in gt14 soil respiration was increased in elevated temperature alone (T) and in elevated O(3) and elevated temperature in combination (O(3) + T) treatments, but in other genotypes O(3) either partly (gt12) or totally nullified (gt25) temperature effects on soil respiration, or acted synergistically with temperature (gt15). Before leaf abscission, all genotypes had the largest leaf biomass in T and O(3) + T treatments, whereas at the end of the season temperature effects on leaf biomass depended on the prevailing O(3) level. Temperature increase thus delayed and O(3) accelerated leaf senescence, and in combination treatment O(3) reduced the temperature effect. Photosynthetic : non-photosynthetic tissue ratios (P : nP ratios) showed that elevated temperature increased foliage biomass relative to woody mass, particularly in gt14 and gt12, whereas O(3) and O(3) + T decreased it most clearly in gt25. O(3)-caused stem growth reductions were clearest in the fastest-growing gt14 and gt25, whereas mycorrhizal root growth and sporocarp production increased under O(3) in all genotypes. A labelling experiment showed that temperature increased tree total biomass and hence (13)C fixation in the foliage and roots and also label return was highest under elevated temperature. Ozone seemed to change tree (13)C allocation, as it decreased foliar (13)C excess amount, simultaneously increasing (13)C excess obtained from the soil. The present results suggest that warming has potential to increase silver birch growth and hence carbon (C) accumulation in tree biomass, but the final magnitude of this C sink strength is partly counteracted by temperature-induced increase in soil respiration rates and simultaneous O(3) stress. Silver birch populations' response to climate change will also largely depend on their genotype composition.     

Interaction between moisture, nutrients and growth of white spruce in interior Alaska

Two thinning and fertilization studies, the first in 1969 and the second in 1971, were established to evaluate the question of nutrient limitation to tree growth and the consequences of stand manipulation of soil moisture supply. Fertilizer was applied yearly for the first 5 years in both studies; growth response has been measured through 1987. Results indicate that thinning is necessary to obtain a growth response to fertilizer applied at the rate of 111 kg nitrogen ha⁻¹. The response to fertilization after fertilization ended lasted for 4 years in plots thinned to 800 stems ha⁻¹, while a significant response continued for only 2 years in plots thinned to 1600 stems ha⁻¹.

A soil water-balance model was calibrated for the control and treatment plots of these two studies. Soil water-deficits were estimated and correlated with yearly average basal-area growth per tree. Results indicated that there is a correlation between seasonal soil-moisture deficit and growth during the years when soil moisture was measured for the unthinned control plots (r² = −0.787, P = 0.002) but not for the thinned and fertilized plots (r² = −0.652, P = 0.057).     

Effects of different soil treatments on growth and net nitrogen uptake of newly planted Picea abies (L.) Karst. seedlings

Five soil treatments in a 4-year-old clearcut in southern Sweden affected biomass increase and net nitrogen uptake by planted Norway spruce (Picea abies (L.) Karst.) seedlings through their on net mineralisation and root growth. The patch soil treatments studied were: (i) soil inversion in an old clearcut; (ii) mineral soil from the clearcut remaining, (iii) mineral soil from the clearcut with fertiliser application during the first season; (iv) mineral soil from a nearby uncut forest replacing the clearcut mineral soil; and (v) an untreated control. Growth increased in seedlings in treatments (i) and (iii), but growth in soil treatments with humus removal was not better than that of seedlings in untreated soil. High N uptake early in the first growing season resulted in increased growth during this season in contrast to late N uptake that resulted in a high N concentration in the seedlings after the first growing season. This in turn led to a high growth rate during the next growing season. Generally, both root growth and net N mineralisation were positively correlated to N uptake in the soil treatments. Therefore, a combination of low net N mineralisation and poor root growth as a result of high soil density appears to explain the low N uptake in seedlings in undisturbed soil. The importance of competition with field vegetation for N and water was not clear. Net mineralisation was larger in soil treatments where the humus layer was retained than where it was removed. Net N mineralisation in soil from old clearcuts was the same as in soil from fresh clearcuts.     

Establishment and growth of white spruce on a boreal forest floodplain: Interactions between microclimate and mammalian herbivory

White spruce (Picea glauca (Moench) Voss) is a dominant species in late-successional ecosystems along the Tanana River, interior Alaska, and the most important commercial timber species in these boreal floodplain forests. Whereas white spruce commonly seed in on young terraces in early primary succession, the species does not become a conspicuous component of the vegetation until after 60–80 years. To address what abiotic and/or biotic factors may explain the paucity of spruce in earlier stages of succession, we examined germination and growth of planted white spruce seedlings across an environmental gradient that included variation in soil physico-chemical properties in the presence and absence of mammal browsing. The effect of browsing pressure over the first four years after planting was most noticeable on the older terraces. Likewise, direct effects of hare browsing on spruce seedling mortality were only manifested at the oldest sites. Spruce germination and survival was inversely proportional to soil cation concentrations, which was largely controlled by temperature-driven evapotranspiration. High light intensities and high air temperatures significantly reduced seedling growth, whereas variation in soil moisture only explained a significant amount of variation in seedling survival. Temperatures within the needle clusters on terminal shoots reached values that adversely affect photosynthesis (>32 °C) on multiple occasions over the growing season. We conclude that the direct (temperature) and indirect (soil chemistry) effects of high insolation are major factors constraining spruce performance on early successional terraces, and that these effects can be significantly exacerbated by mammal browsing on associated deciduous vegetation.     

The effects of sulfometuron on the root growth of loblolly pine

The effect of sulfometuron (Oust) on the root growth of loblolly pine (Pinus taeda L.) seedlings was studied in the field and in two soil types in the greenhouse. In the greenhouse study sulfometuron was applied to the foliage only, the soil only, and to both foliage and soil at the rates of 0.10, 0.21, and 0.42 kg ai/ha. Twenty-eight days after application root growth significantly decreased for all methods of application and with increasing rates e.g., 0.10, 0.21, and 0.42 kg ai/ha sulfometuron reduced new root length by 42%, 53%, and 64%, respectively. Application in the field at the rate of 0.30 kg ai/ha resulted in a 68% reduction in root growth. However, by the end of the growing season treated seedling root and shoot biomass were 115% and 64% greater, respectively. To ensure earlier season survival the rate of sulfometuron applied should be kept at a minimum level that will allow both weed control and adequate root growth. Low rates would be particularly important during a dry planting season.     

Seasonal shoot and needle growth of loblolly pine responds to thinning, fertilization, and crown position

The impacts of thinning, fertilization and crown position on seasonal growth of current-year shoots and foliage were studied in a 13-year-old loblolly pine (Pinus taeda L.) plantation in the sixth post-treatment year (1994). Length of new flushes, and their needle length, leaf area, and oven-dry weight were measured in the upper and lower crown from March through November. Total shoot length was the cumulative length of all flushes on a given shoot and total leaf area per shoot was the sum of leaf areas of the flushes.

By the end of June, first-flush foliage reached 70% of the November needle length (14.3 cm) and 65% of the final leaf area (15.0 cm²). Cumulative shoot length of first- and second-flush shoots achieved 95% of the annual length (30.3 cm), whereas total leaf area per shoot was 55% of the final value (75.3 dm²). Fertilization consistently stimulated fascicle needle length, dry weight, and leaf area in the upper crown. Mean leaf area of upper-crown shoots was increased by 64% six years after fertilization. A significant thinning effect was found to decrease mean leaf area per shoot in the crown. For most of the growing season, the thinned-fertilized trees produced substantially more leaf area per shoot throughout the crown than the thinned-nonfertilized trees. These thinned-fertilized trees also had greater needle length and dry weight, longer first flush shoots, and more leaf area per flush than trees in the thinned-nonfertilized plots. Needle length and leaf area of first flush shoots between April and July were linearly related to previous-month canopy air temperature (T_a). Total shoot length strongly depended on vertical light gradient (PPFD) within the canopy, whereas shoot leaf area was a function of both PPFD and T_a. Thus, trees produced larger and heavier fascicles, more and longer flush shoots, and more leaf area per shoot in the upper crown than the lower crown. We conclude that thinning, fertilization, and crown position regulate annual leaf area production of current-year shoots largely by affecting the expansion of first flush shoots and their foliage during the first half of the growing season.     

Willow stand growth model based on nitrogen productivity

A simple model for willow forest growth during the growing season is presented. The model is based on nitrogen productivity, expressed as the daily above‐ground biomass growth per the amounts of nitrogen in foliage and leaf litter. It is shown that the nitrogen productivity can be expressed as a linear function of the amounts of leaves produced. The model, which also includes allocation of biomass between stem and leaf biomass as well as between foliage and leaf litter biomass, is tested against accurate measurements of the seasonal growth of willow of four different plots. Simulated and measured biomass of stem, foliage, leaf litter and the total above‐ground biomass agreed well for all four plots. The model concept should be useful in nitrogen flux models and as a tool to describe the actual growth of a willow stand, knowing only the annual leaf production and the nitrogen concentrations of foliage during the growing season.     

Microclimate and soil moisture effects of three intercrops on the rows of a newly-planted intercropped plantation

Trees are grown in intercropping systems for a variety of purposes including wood products, fuelwood, fruit, forage or conservation purposes. No matter what end use, different tree/crop combinations interact differently resulting in differential growth rates of the trees during establishment. Preliminary work has shown that seedling growth and survival of trees are related to their intercrop and the results of this study help to explain these findings. Soil water potential, soil and air temperature, relative humidity, windspeed, and light (photosynthetic photon flux density — PPFD) were measured throughout the growing season in the clean-weeded treerows within crops of corn, soybeans and winter wheat. Crop height and biomass were also measured. This study was conducted during the 1992 growing season which was unusually cool and wet from mid June into the winter. The growth of winter wheat, measured by crop height and above-ground biomass, was earlier in the season than that of soybeans and corn, and this pattern affected the environmental conditions in the tree rows. Soil water potential was affected with associated effects on soil temperature (in combination with other factors). Crop height drastically reduced windspeed in the corn treatment from July through winter, also affecting PPFD and soil temperature later in the year. Although many microclimate differences were relatively small, data from subsequent years as well as associated soil moisture studies and additional years will help to further elucidate these relationships.     

Biomass and carbon sequestration of ponderosa pine plantations and native cypress forests in northwest Patagonia

Fast growth tree plantations and secondary forests are considered highly efficient carbon sinks. In northwest Patagonia, more than 2 million ha of rangelands are suitable for forestry, and tree plantation or native forest restoration could largely contribute to climate change mitigation. The commonest baseline is the heavily grazed gramineous steppe of Festuca pallescens (St. Yves) Parodi. To assess the carbon sequestration potential of ponderosa pine (Pinus ponderosa (Dougl.) Laws) plantations and native cypress (Austrocedrus chilensis (Don) Flor. et Boutl.), individual above and below ground biomass models were developed, and scaled to stand level in forests between 600 and 1500 annual rainfall. To calculate the carbon sequestration baseline, the pasture biomass was simulated. Also, soil carbon at two depths was assessed in paired pine-cypress-pasture sample plots, the same as the litter carbon content of both forest types. Individual stem, foliage, branch and root log linear equations adjusted for pine and cypress trees presented similar slopes (P>0.05), although some differed in the elevations. Biomass carbon was 52.3 Mg ha⁻¹ (S.D.=30.6) for pine stands and 73.2 Mg ha⁻¹ (S.D.=95.4) for cypress forests, given stand volumes of 148.1 and 168.4 m³ ha⁻¹, respectively. Soil carbon (litter included) was 86.3 Mg ha⁻¹ (S.D.=46.5) for pine stands and 116.5 Mg ha⁻¹ (S.D.=38.5) for cypress. Root/shoot ratio was 19.5 and 11.4%, respectively. The low r/s value for cypress may account for differences in nutrient cycling and water uptake potential. At stand level, differences in foliage, taproot and soil carbon compartments were highly significative (P<0.01) between both forest types. In pine stands, both biomass and soil carbon were highly explained by the rainfall gradient (r²=0.94). Nevertheless, such a relationship was not found for cypress, possibly due to stand and soil disturbances in sample plots. The carbon baseline estimated in pasture biomass, including litter, was 2.6 Mg ha⁻¹ (S.D.=0.8). Since no differences in soil carbon were found between pasture and both forest types, additionality should be accounted only by biomass. However, the replacement of pasture by pine plantations may decrease the soil carbon storage, at least during the first years. On the other hand, the soil may be a more relevant compartment of sequestered carbon in cypress forests, and if pine plantation replaces cypress forests, soil carbon losses could cause a negative balance.     

Effects of soil moisture and species composition on growth and productivity of trembling aspen and white spruce in planted mixtures: 5-year results

Aspen (Populus tremuloides Michx.) and white spruce (Picea glauca (Moench.) Voss) were planted 0.5 m apart in intimate mixtures in 5 × 4 m plots, with two moisture regimes—irrigation versus control—and five species compositions—pure aspen (Aw100), mixed aspen and spruce (Aw83Sw17, Aw50Sw50, Aw17Sw83), and pure spruce (Sw100), replicated six times. Fifth-year assessments indicated that irrigation increased individual tree growth (height, RCD, crown width), plot leaf area index (LAI), and wood biomass. Increased aspen composition reduced the availability of soil moisture and consequently the growth of individual trees. With increased aspen composition more growth was allocated to stem in aspen and to foliage in white spruce. Comparatively, aspen responded more to irrigation and thus their growth is more dependent on precipitation than that of spruce. Among the three growth variables assessed, height responded more to irrigation in both species. Equal mixtures and aspen-dominated mixtures in control plots had higher productivity in terms of total wood biomass in both absolute and relative terms. The implications of these findings are discussed in relation to managing aspen and white spruce mixedwood forests under increasing drought expected as a result of climate change.     

Pipe-model ratio distributions and branch foliage biomass: differences between two sympatric spruce species

The foliage biomass–sapwood relationship (the pipe model) is critical for tree growth and is used in tree growth models for understanding the implications of this structural relationship on the allocation of resources. In this research, we compared this relationship for two commercially important and sympatric species, black spruce (Picea mariana (Mill.) B.S.P.) and white spruce (Picea glauca (Moench) Voss). At locations in eastern Canada, 57 black and 50 white spruce trees were destructively sampled to obtain foliage biomass, crown structure, and tree stem measures. Using a model-based approach, we compared foliage biomass–branch basal area and foliage biomass–sapwood relationships at the tree and disk (i.e. along the tree stem) levels (i.e. pipe-model ratios) between these two species. We found that (i) branch foliage biomass–branch basal area was greater for black spruce than white spruce and (ii) pipe-model ratios along the tree stem given tree size were greater for black spruce than for white spruce. We attributed these differences to: (i) greater shade tolerance and leaf longevity of black spruce; (ii) slower growth rates of black spruce; and (iii) differing hydraulic strategies and mechanical requirements.     

Influence of woody and herbaceous competition on microclimate and growth of eastern white pine (Pinus strobus L.) seedlings planted in a central Ontario clearcut

The influence of woody and herbaceous plant competition, either alone or in combination, on microclimate and growth of planted eastern white pine (Pinus strobus L.) seedlings was examined over four consecutive growing seasons in a central Ontario clearcut. Treatments that manipulated the comparative abundance of these two plant functional groups significantly affected light availability, soil moisture, and air and soil temperature regimes. These microclimate alterations, coupled with the relative competitiveness of herbaceous and woody vegetation, corresponded to temporal changes in vegetation cover and dominance. The more rapid colonization and growth of the herbaceous plant community, dominated by bracken fern (Pteridium aquilinum) and ericaceous shrubs (Kalmia sp., Vaccinium sp.), resulted in this form of vegetation being a comparatively important early competitor for soil moisture. As the woody plant community, dominated by naturally regenerated trembling aspen (Populus tremuloides Michx.), grew in height and leaf area, it became a comparatively strong competitor for both light and soil moisture. For all vegetation treatments combined, white pine seedling growth responses were strongly correlated with total cover of competing vegetation and its relative influence on above- and belowground microclimatic variables. Higher total cover of competing vegetation was generally associated with lower light and soil moisture availability and cooler soil temperatures. Multiple regression analyses indicated that pine seedling relative height growth increased with soil moisture content and growing season soil heat sum, while seedling relative diameter and relative volume growth increased with light availability.     

Growth, nutrition, photosynthesis and transpiration responses of longleaf pine seedlings to light, water and nitrogen

Early growth and physiology of longleaf pine (Pinus palustris Mill.) seedlings were studied in response to light, water and nitrogen under greenhouse conditions. The experiment was conducted with 1-year-old seedlings grown in 11.3 l pots. The experimental design was a split-plot factorial with two levels (low and high) of each of the factors, replicated in three blocks. The four factorial combinations of water and nitrogen were randomly applied to 15 pots (sub-plots) in each of the light treatment (main plot). Data were collected on survival, root collar diameter (RCD), and height on a monthly basis. Biomass (shoot, root and needle), leaf area index, specific needle area, and needle nutrient (N, P, K, Ca, and Mg) concentrations were determined following final harvest after 16 months. Physiological data (net photosynthesis and transpiration) were collected monthly from March to July during the second growing season.

Height and RCD were significantly influenced by nitrogen and water and by the interaction between them with no apparent effect of light. Seedlings grew 93% taller in the high nitrogen and well watered (HNWW) treatment compared to the low nitrogen and water stressed (LNWS) treatment. Similarly, a significant increase (78%) in RCD was observed for seedlings in the HNWW treatment over the LNWS treatment. Light, along with water and nitrogen, played an important role in seedling biomass growth, especially when water was not limiting. Biomass partitioning (as measured by root:shoot ratio) was affected only by nitrogen and water. Nutrient stress had a greater influence on carbon allocation (69% increase in root:shoot ratio) than water stress (19% increase). Net photosynthesis (P_net) was significantly higher for seedlings in the high resource than in the low resource treatments with significant light×water and nitrogen×water interactions. Transpiration rate was higher (75%) under the WW treatment compared to the WS treatment. Longleaf pine seedlings grown under the LNWW treatment had the lowest foliar nitrogen (0.71%) whereas seedlings in the HNWS treatment had the highest (1.46%). Increasing the availability of light (through larger canopy openings or controlling midstory density) and soil nitrogen (through fertilization) may not result in greater P_net and improved seedling growth unless soil water is not limiting.     

Ailanthus triphysa at different densities and fertiliser regimes in Kerala, India: growth, yield, nutrient use efficiency and nutrient export through harvest

A split plot trial involving Ailanthus triphysa (ailanthus) at four spacings (3 m×1 m, 2 m×2 m, 3 m×2 m and 3 m×3 m) and four fertiliser regimes (0:0:0, 50:25:25, 100:50:50 and 150:75:75 kg ha⁻¹ per year N, P₂O₅, K₂O) was initiated in June 1991. Objectives included evaluating the growth and yield potential of ailanthus grown under differing density and fertiliser regimes and to estimate the nutrient export through harvest. Ninety-six randomly selected average-sized trees were felled at 8.8 years of age for assessment. Results show that height, diameter, stand leaf area index, biomass production and volume yield were greater in the 2 m×2 m spacing. Repeated application of fertilisers at 1.2, 2.25 and 5.25 years after planting had little effect on biomass and volume yields, presumably because of weed competition (despite periodic weed control), higher pest incidence (in the heavily fertilised plots) and/or moderately adequate soil nutrient levels. Regarding partitioning of tree biomass, stem wood represented the principal component (>70%), while foliage contributed the least (<7%). Conversely, foliar N, P and K concentrations were the highest, followed by branch wood, coarse roots and stem wood. Denser stands showed greater accumulation of N, P and K with higher potential for nutrient export through harvest. However, as the bole fraction accounted for only about 56–64% of the total nutrients removed, leaving other biomass components (foliage and branches) at the site will reduce the associated nutrient export. Wider spacings (3 m×2 m and 3 m×3 m) were more efficient in N and K use, but P use efficiency was higher in 2 m×2 m. Likewise, trees in the no fertiliser plots exhibited greatest N, P and K use efficiencies. Available soil P, K and organic C levels declined with increasing tree density, while repeated fertilisation increased nutrient concentrations. Soil pH and available P levels declined in comparison to the pre-treatment values.