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.     

Inhibition of protein synthesis in loblolly pine hypocotyls by mannitol-induced water stress

The relationship between mannitol-induced water stress and protein synthesis was investigated in hypocotyl slices of loblolly pine (Pinus taeda L.). Mannitol-induced water stress inhibited the incorporation of L-[(35)S]methionine into protein. As the water potential decreased, incorporation of label into protein decreased in both the soluble- and membrane-protein fractions. There were no significant differences in response to water stress among seed sources from four different geographical regions.     

Acquired thermotolerance of jack pine, white spruce and black spruce seedlings

The acquired thermotolerance of first-year seedlings of jack pine (Pinus banksiana Lamb.) hardened at 36, 38, 40 or 42 degrees C for 90, 180 or 360 minutes and of black spruce (Picea mariana (Mill.) B.S.P.) hardened at 34, 36, 38 or 40 degrees C for 30, 90, 180 or 360 minutes was determined by comparison of needle damage to that of non-hardened seedlings (25 degrees C) following exposure to temperatures of 49 and 47.5 degrees C, respectively. Compared to seedlings kept at 25 degrees C, heat injury sustained from exposure to high temperatures was markedly reduced following hardening for 180 minutes at 36 and 38 degrees C in jack pine and black spruce, respectively. Increasing the exposure time at 36 degrees C in jack pine, and at 36 to 40 degrees C in black spruce, also reduced needle damage. The duration of increased thermotolerance was investigated in jack pine, black spruce and white spruce (Picea glauca (Moench) Voss) by comparing heat injury from high temperatures in non-hardened seedlings and in seedlings hardened at 38 degrees C for 180 minutes a day for either 1, 3 or 6 days. In all three species, the duration of acquired thermotolerance increased with the number of days of heat hardening. For jack pine and white spruce seedlings hardened at 38 degrees C for 6 days, increased thermotolerance persisted for at least 14 and 10 days, respectively, after the end of the hardening treatment. In contrast, the thermotolerance of black spruce seedlings hardened at 38 degrees C for 6 days remained elevated for only 4 days.     

Reduction in turgid water volume in jack pine,white spruce and black spruce in response to drought and paclobutrazol

Significant reductions in needle water content were observed in white spruce (Picea glauca (Moench) Voss), black spruce (Picea mariana (Mill) B.S.P.), and jack pine (Pinus banksiana Lamb.) seedlings in response to a 10-day drought, although turgor was apparently maintained. When the seedlings were re-watered after the drought, jack pine needles regained their original saturated volume, whereas white spruce and black spruce needles did not. Significant drought-induced reductions in turgor-loss volume (i.e., tissue volume at the point of turgor loss) were observed in shoots of all three species, especially jack pine. Repeated exposure to 7 days of drought or treatment with the cytochrome P(450) inhibitor, paclobutrazol ((2RS,3RS)-1-(4-chlorophenyl)-4,4-dimethyl-2-(1,2,4-triazol-1-yl)-pentan-3-ol), reduced seedling height relative to that of untreated controls in all three species. The reductions in saturated and turgor-loss needle volumes in the paclobutrazol-treated seedlings were comparable with those of seedlings subjected to a 10-day drought. The treatment-induced reductions in shoot and needle water contents enabled seedlings to maintain turgor with tissue volumes close to, or below, the turgor-loss volume of untreated seedlings. Paclobutrazol-treated seedlings subsequently survived drought treatments that were lethal to untreated seedlings.     

Water relations and morphological development of bare-root jack pine and white spruce seedlings: seedling establishment on a boreal cut-over site

Bare-root jack pine (Pinus banksiana Lamb.) and white spruce (Picea glauca (Moench) Voss) seedlings were planted on a boreal cut-over site and subsequent growth and seedling water relation patterns were monitored over the first growing season. Comparison of morphological development between white spruce and jack pine showed jack pine seedlings had greater new root development and a lower new shoot/new root ratio, while white spruce seedlings had greater new shoot development. Seasonal water relation patterns showed white spruce seedlings to have a greater decrease in xylem pressure potential (_x) per unit increase in transpirational flux density in comparison to jack pine seedlings. These results suggest that the greater resistance to water flow through the soil-plant-atmosphere continuum in white spruce seedlings compared to jack pine seedlings may be due to the relative lack of new root development in white spruce. Stomatal response of the seedlings showed that as absolute humidity deficit between the needles and air (AHD) increased, needle conductance (gwv) decreased in both species, but at low AHD levels white spruce had gwv approximately 35% higher than jack pine. For white spruce seedlings, gwv decreased as _x became more negative in a predictable curvilinear manner, while gwv of jack pine seedlings responded to _x with a threshold closure phenomenon at approximately - 1.75 MPa. Tissue water potential components for jack pine and white spruce seedlings at the beginning and end of the growing season showed jack pine to reach turgor loss at 76% relative water content while white spruce reached turgor loss at 88% relative water content. White spruce seedlings showed osmotic adjustment over the growing season, with an osmotic potential at turgor loss of - 1.27 MPa and - 1.92 MPa at the beginning and end of the growing season, respectively. Jack pine did not show any osmotic adjustment over the growing season. The implication of morphological development on water relation patterns are discussed with reference to successful seedling establishment.     

Winter water relations of New England conifers and factors influencing their upper elevational limits. I. Measurements

The upper elevational limits of tree species are thought to be controlled by abiotic factors such as temperature and the soil and atmospheric conditions affecting plant water status. We measured relative water contents (RWC), water potentials (Psi) and cuticular conductances (g(c)) of shoots of four conifer species-eastern hemlock (Tsuga canadensis (L.) Carr.), eastern white pine (Pinus strobus L.), red pine (P. resinosa Ait.) and red spruce (Picea rubens Sarg.)-during two winters on Mt. Ascutney, Vermont, USA. Some micrometeorological measurements are also reported. Eastern hemlock and white pine were studied near their upper elevational limits at a 640-m site, and red pine was studied near its upper elevational limit at 715 m. Red spruce was also studied at the 715-m site, which is in the middle of its elevational range on this mountain. There was no evidence of winter desiccation stress in any species. The observed distribution of seedlings suggested that the upper elevational limits on shade-intolerant eastern white pine and red pine are set by the absence of suitable seed beds after 100 years without fire. Eastern hemlock is able to reproduce in deep shade on organic substrates, but germination at high elevations may be restricted by low temperatures.     

Effects of near-lethal heat stress on bud break, heat-shock proteins and ubiquitin in dormant poplar (Populus nigra Charkowiensis x P. nigra incrassata)

We assessed the effects of near-lethal heat stress on bud break, heat-shock proteins (HSPs) and ubiquitin in hybrid poplar (Populus nigra (L.) Charkowiensis x P. nigra (L.) incrassata). Shoots, with 10-15 buds each, were collected from September to March and exposed to temperatures between 20 and 60 degrees C for 2 h. Shoots were then placed in a greenhouse at 18-22 degrees C with supplemental light and cumulative bud break was recorded over a 4-week period. Samples of bud tissues were collected during and up to 96 h after heat treatment for protein analysis. De novo synthesis of proteins was monitored by exposing excised buds to [(35)S]-methionine for 3 h before, during, or after heat treatment. Heat treatments of 40-45 degrees C resulted in both a release from endodormancy and a decrease in thermal units needed for bud break during ecodormancy. The response to near-lethal heat stress was complex and was affected by intrinsic thermal sensitivity. Heat treatments were least effective during August and became progressively more effective as endodormancy progressed. In the later stages of ecodormancy, a heat treatment of 45 degrees C either inhibited bud break or killed the buds. Although temperatures of 42.5 to 45 degrees C inhibited incorporation of [(35)S]-methionine into proteins for at least 48 h, several HSPs were synthesized in response to temperatures of 40-45 degrees C. Immunoblots indicated that one of the heat-induced proteins was immunologically related to HSP70. Increases in free and conjugated forms of ubiquitin were also observed in response to heat treatment. Production of HSPs and ubiquitin, however, was not consistently associated with the heat treatments that induced the highest percentage of bud break. The roles of heat-induced protein degradation, HSPs, and ubiquitin in overcoming dormancy by near-lethal heat stress are discussed.     

Responses of Pinus sylvestris and Picea abies Seedlings to Limited Phosphorus Fertilization and Treatment with Elevated Ozone Concentrations

Scots pine ( Pinus sylvestris L.) and Norway spruce [ Picea abies (L.) Karst.] seedlings were exposed to high phosphorus (HP) or low phosphorus (LP) availability for one growing season in the open field, and to combined P availability and elevated ozone (O 3 ) concentrations (0, 55, 110 and 210 ppb for Scots pine and 0, 40, 75 and 150 ppb for Norway spruce, respectively) for 28 days in controlled laboratory chambers. Compared with HP, the LP treatment reduced Scots pine current-year (C) shoot and root dry masses and Norway spruce total dry mass, whereas the highest O 3 concentrations increased the magnesium concentration of Scots pine C needles and P concentrations of the C needles of both tree species. Chlorophyll a, a+b and carotenoid concentrations of Scots pine C needles were significantly higher in the LP treatment compared with HP under the highest O 3 concentration (210 ppb). In the mesophyll tissue of C needles of both tree species, LP treatment increased the size of mitochondria and elevated O 3 -induced granulation of chloroplast stroma and disintegration of cytoplasm. Exposure to elevated O 3 concentrations increased swelling of chloroplast thylakoids and reduced the amount of vacuolar tannin in the LP Scots pine C needles. The results suggest disturbances in needle photosynthetic machinery due to acute exposure to the combination of elevated O 3 and low P availability. However, clear additive effects were found only in needle P concentrations < 1 mg g -1 in short-term O 3 exposure.     

Damage by pine weevil Hylobius abietis to conifer seedlings after shelterwood removal

Abstract

Pine weevil (Hylobius abietis L.) damage to seedlings after overstorey removal was investigated in a survey study in six shelterwoods in the south–central part of Sweden. The shelterwoods predominantly consisted of Scots pine, except at one site where the shelter trees mainly consisted of Norway spruce. Before final cutting, 10 plots were laid out at each site and measurements of shelter trees and marked seedlings were taken. The seedlings were examined during the 2 years after final cutting. The study showed that removal of shelter trees increases the risk of severe damage by pine weevil and the variable that was most strongly correlated with the risk was the seedling root collar diameter. Both Scots pine and Norway spruce seedlings were severely damaged by pine weevil, and most of the feeding occurred during the first year after cutting. The amount of debarked area was significantly larger for Scots pine than for Norway spruce seedlings. Vitality (growth of the leading shoot before final cutting) of the seedlings also affected the probability of damage. Seedlings with high vitality were less damaged by pine weevil than seedlings with low vitality. For Scots pine the shelterwood density before final cutting was correlated to the intensity of pine weevil feeding after cutting. In conclusion, after the final cutting of a pine or spruce shelterwood, pine weevils will probably invade the area. To avoid serious damage, Norway spruce and Scots pine seedlings should have reached a diameter of at least 10–12 mm.     

Interference of bracken (Pteridium aquilinum L. Kuhn) with Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies L. Karst.) seedling establishment

The nature of interference of bracken with Scots pine and Norway spruce seedling establishment was considered in three field experiments. In a seeding experiment, it was found that Scots pine germination was highest on exposed mineral soil and lowest when intact bracken litter and humus were present, suggesting adverse effects of litter and humus on pine regeneration probably due to phytotoxicity. In a second experiment, smothering by bracken caused high mortality of Scots pine seedlings while Norway spruce seedlings were relatively unaffected. Mortality for both Scots pine and Norway spruce seedlings was low when planted in a adjacent Scots pine-bilberry stand with no bracken. Annual shoot growth of Norway spruce was higher in bracken than in Scots pine-bilberry vegetation while no differences in shoot growth between these two vegetation types occurred for Scots pine. In a third experiment, activated carbon was added to the ground under Norway spruce seedlings planted in bracken to adsorb possible phytotoxic compounds released by bracken. The addition of carbon had no effect on seedling mortality or growth rate, indicating that the seedlings were not susceptible to allelochemicals released by bracken. Since large Norway spruce seedlings were relatively unaffected by bracken interference in this study, artificial regeneration with containerized Norway spruce seedlings is suggested to achieve an acceptable conifer tree establishment on clear-cuts invaded by bracken.     

Morphological characteristics associated with tolerance to competition from herbaceous vegetation for seedlings of jack pine,black spruce,and white pine

Tolerance of bareroot and container-grown seedlings of black spruce (Picea mariana (Mill.) B.S.P.), jack pine (Pinus banksiana Lamb.), and eastern white pine (Pinus strobus L.) to competition from herbaceous vegetation was examined in the first five years after planting on a site in the Great Lakes/St. Lawrence forest of Ontario, Canada. Shoot and root morphological characteristics of various stocktypes were measured before planting and correlated with 5-year survival and growth following control and no control of herbaceous vegetation. For black spruce and jack pine, medium-sized bareroot stocktypes had greater relative 5-year stem volume growth in the presence of herbaceous vegetation than did container stock of either species or large bareroot stock of spruce. Relative volume growth was measured as the ratio of the cumulative stem volume increment in the presence of vegetation (Veg) to that in the absence of vegetation (NoVeg), i.e., the Veg:NoVeg ratio. In white pine, the Veg:NoVeg ratio of volume increment of medium container and large bareroot stocktypes exceeded that of small container and medium bareroot stocktypes. In jack pine, root collar diameter at planting and number of first-order lateral roots were positively correlated with 5-year Veg:NoVeg ratio of volume increment. In white pine, the Veg:NoVeg ratio was also positively correlated with root collar diameter at planting and with root volume. In black spruce, the ratio was not related to pre-plant morphology. Thus, for white pine and jack pine, certain pre-plant morphological features may be useful in forecasting the relative ability of different stocktypes to grow under herbaceous competition conditions in the field.     

Infection experiments with Gremmeniella abietina on seedlings of Norway spruce and Scots pine

Conidia of Gremmeniella abietina infected and caused disease symptoms in annual shoots of both Scots pine (Pinus sylvestris) and Norway spruce (Picea abies) seedlings. In Norway spruce shoots the infection remained largely latent, with only a few seedlings showing symptoms. Mycelial growth inside the shoots was faster in Scots pine than in Norway spruce and was favoured by low temperature in both hosts. The shoots of Norway spruce seedlings had higher endophyte populations than those of Scots pine, and the populations were decreased by low temperatures. Reductions in the normal epiphytic or endophytic flora by acid mist treatments seemed to favour the development of G. abietina.     

Effects of root temperature on growth and photosynthesis in conifer seedlings during shoot elongation

Growth and gas exchange characteristics were studied in pine (Pinus sylvestris L.) and spruce (Picea abies Karst.) seedlings grown in hydroponic culture in the presence of N (50 mg l(-1)) and transferred at the start of their second growing season to tap water at 5, 8, 12, 16 or 20 degrees C (air temperature between 18-20 degrees C) for 3 weeks (pine) or 5 weeks (spruce). Root growth of both species was completely inhibited at root temperatures of 5 and 8 degrees C, but increased almost exponentially as root temperature increased. Shoot growth was maximal at 12 degrees C in both pine and spruce and decreased at low root temperatures. In both species, CO(2) uptake was decreased at low root temperatures and appeared to be influenced by the pattern of nitrogen retranslocation. In pine seedlings, as root temperature increased, an increasing proportion of the total nitrogen pool was retranslocated to the new shoot, whereas in spruce seedlings nitrogen was retranslocated to the roots. Differences in the retranslocation of nitrogen in the two species were reflected in the amount of soluble protein in needles, which at the end of the experiment increased with increasing root temperature in pine, but decreased in spruce. Our data suggest that in spruce, but not pine, CO(2) uptake was limited by the amount of Rubisco.     

Selection of white spruce families in the context of climate change: heat tolerance

To assess the responses and plasticity of white spruce seedlings (Picea glauca (Moench) Voss) to high temperatures, 12 open-pollinated families differing in growth performance were exposed to a 30-min heat treatment of 42, 44, 46, 48, or 50 degrees C with or without heat preconditioning at 38 degrees C for 5 h. Damage was evaluated based on chlorophyll fluorescence parameters after heat preconditioning, after the heat treatments and during a 7-day recovery period. Visible needle damage was also evaluated after the heat treatments and 14 days later. Chlorophyll fluorescence parameters indicated that seedlings subjected to a heat treatment of 42-43 degrees C lost the ability to phosphorylate and donate water to photosystem II (PSII). A heat treatment of 44-46 degrees C severely limited the ability of the seedlings to use NADPH and ATP in the Calvin cycle. Based on visible needle damage, families with superior height-growth performance were more sensitive to heat stress than families with intermediate or inferior height-growth performance. Moreover, families with superior height-growth performance had low photochemical efficiencies in the light (DeltaF/F(m)') after heat treatment. Heat preconditioning increased the thermotolerance of the seedlings. However, the data suggest that white spruce seedlings exhibiting fast-growing characteristics under present conditions may not grow as well at higher temperatures.     

多重复干旱循环对北美短叶松和黑云杉苗木气体交换及水分利用效率的影响

本文研究了多重复干旱循环对１年生北美短叶松（ＰｉｎｕｓｂａｎｋｓｉａｎａＬａｍｂ．）和黑云杉（Ｐｉｃｅａｍａｒｉａｎａ［Ｍｉｌ］Ｂ．Ｓ．Ｄ．）苗木的气体交换速率及水分利用效率的影响。结果表面，多重干旱循环对它们的气体交换（Ｃｓ，Ｐｎ，Ｔｒ）有显著影响（Ｐ＜０．５），而对其水分利用效率（ＷＵＥ）影响不大（Ｐ＞０．１）。尽管北美短叶松的气孔对轻度干旱胁迫不如黑云杉敏感，但是它对中度及严重干旱胁迫的敏感程度却高于黑云杉。在轻度及中度干旱胁迫下，北美短叶松的光合作用主要受非气孔因素的影响，而黑云杉则主要受气孔因素的影响。解除干旱胁迫后，黑云杉的气孔敏感性、光合能力及水分利用效率的恢复都要比北美短叶松更快．我们认为，延迟脱水是北美短叶松的主要耐旱机理，而忍耐脱水则是黑云杉重要的耐旱途径。轻度的干旱胁迫锻炼可以帮助北美短叶松在更严重的干旱胁迫下保持固有而较强的耐旱能力。然而，通过多重复干旱循环锻炼后黑云杉在改善耐旱能力的强度方面则大于北美短叶松     

CONIFEROUS LITTER AMENDMENTS AND THE GROWTH OF SITKA SPRUCE

The growth of semi-checked Sitka spruce (Picea sitcbensis Carr.)on heathland has been stimulated to different degrees by theannual application of litter of Scots pine (Pinus sylvestrisL.), Corsican pine (P. nigra var. calabrica Schneid), Lodgepolepine (P. contorta Dougl.), Japanese larch (Larix leptolepisGord.), and Sitka spruce, at rates corresponding to normal plantationconditions, following an initial heavier rate. Foliar analysissuggests that the response is at least partially attributableto the influence of the litters on the nitrogen nutrition ofthe spruce and differences between litters appear to be largelydetermined by their total nitrogen contents. In this respect,Japanese larch, with an annual needle fall generally greaterthan that of the pines and a higher nitrogen content, offersthe best promise as a nurse species for spruce.     

Changes in protein synthesis during drought conditioning in roots of jack pine seedlings (Pinus banksiana Lamb.)

The impact of drought conditioning on the ability of eight-week-old jack pine (Pinus banksiana Lamb.) seedlings to withstand drought was assessed. Two progressive cycles of drought conditioning significantly increased the survival of seedlings subjected to a subsequent prolonged drought. The in vivo accumulation of several root membrane proteins during drought conditioning was correlated with an increase in seedling survival. A group of root proteins, ranging in molecular mass from 43 to 47 kDa, increased accumulation during one cycle of drought conditioning and to a lesser extent during two cycles of drought conditioning. The accumulation of several low molecular mass membrane and soluble proteins also increased during drought conditioning, suggesting that these proteins may play an important role in the enhancement of drought tolerance. In vitro translation studies showed a general increase in the abundance of protein products encoded by mRNAs from drought-conditioned seedlings. Although the majority of the in vitro translation products appeared in both control and drought-conditioned seedlings, one mRNA encoding a 15 kDA translated protein was more prominent during the second cycle of drought conditioning.     

Infection of Norway spruce container seedlings by Gremmeniella abietina

First‐ and second‐year containerized Norway spruce seedlings were inoculated with conidia of type A (large tree type) and type B (small tree type) of Gremmeniella abietina var. abietina at different times during the summer. The appearance of symptoms after artificial inoculation and natural infection on spruce seedlings were recorded the following spring and compared with the disease symptoms on Scots pine seedlings. The proportion of diseased seedlings after inoculation reached as high as 80%. The susceptible period during the summer began later on the first‐year seedlings than on the second‐year seedlings, and was similar for the pine seedlings. Susceptibility of first‐year seedlings was highest in August and on second‐year seedlings in July. The accumulated temperature sum, relative humidity and height growth for first‐ and second‐year seedlings was assessed. Natural infection in 2002 caused more disease on pine than on spruce seedlings. Experimental thinning of seedlings had no effect on disease incidence. In a preliminary comparison between the ability of A and B types to cause disease in Norway spruce seedlings, type B caused more damage than type A after inoculation. However, type A caused a high disease frequency in other experiments in this study. Symptoms on Norway spruce seedlings often first occurred in the mid‐section of the shoot, and were similar to those observed on pine seedlings: needles turned brown, starting at the needle base, in the spring following inoculation. On first‐year spruce, diseased needles were shed rapidly, in contrast to a slower rate of shedding on first‐year pine seedlings. Pycnidia developed about 2 years after inoculation (on pine 1 year after inoculation). On Norway spruce seedlings the lower part of the shoot, including the lateral shoots, often remained alive. The experiments show that G. abietina can cause disease on containerized Norway spruce seedlings under nursery conditions in Finland. The coincidence of spore dispersal, seedling susceptibility and predisposing factors are important in disease development.     

Height growth of planted conifer seedlings in relation to solar radiation and position in Scots pine shelterwood

Seedlings of different provenances of Scots pine (Pinus sylvestris L.), lodgepole pine (Pinus contorta Dougl., var. latifolia Engelm.) and Norway spruce (Picea abies (L.) Karst.) were planted in three Scots pine shelterwoods (125, 65 and 43 stems ha⁻¹) and a clear-cut, all in northern Sweden. The sites were mounded and planting took place during 2 consecutive years (1988 and 1989). The solar radiation experienced by the individual seedlings was determined using a simulation model. Height development of the seedlings was examined during their first 6 years after planting. During the final 3 years of the study, height growth of Norway spruce was relatively poor, both in the shelterwoods and the clear-cut area. Height growth of lodgepole pine was significantly greater than that of Scots pine, both in the shelterwoods and the clear-cut. In contrast to Norway spruce, Scots pine and lodgepole pine displayed significantly greater height growth in the clear-cut than in the shelterwoods. For all three species in the shelterwoods, regression analyses showed that height growth was more strongly correlated with the distance to the nearest tree than with the amount of radiation reaching the ground, i.e. growth was reduced in the vicinity of shelter trees. Therefore, we conclude that the significant reduction in height growth of seedlings of Scots pine and lodgepole pine in Scots pine shelterwoods was partially caused by factors associated with the distance to the nearest shelter tree. Because the substrate was a nitrogen-poor sandy soil, we suggest that root competition for mineral nutrients, especially nitrogen, accounts for the reduction in height growth.