Root growth capacity and field performance of Pinus sylvestris and Picea abies seedlings

Correlations between root growth capacity (RGC), at the time of planting, and field performance were studied for Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) seedlings. Before planting a gradient in seedling viability was generated through exposure to low root temperatures and different winter storage regimes. The hypothesis that high RGC values would improve field performance was to some extent verified for pine seedlings while no correlations could be registered for spruce. Reasons for these results are discussed.     

Freezing temperatures in the root zone—effects on growth of containerized Pinus sylvestris and Picea abies seedlings

Roots of 1‐year‐old containerized seedlings of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) were experimentally frozen in December. The seedlings were then grown for 3 weeks in a growth chamber and evaluated with regard to root growth capacity (RGC) and shoot elongation. The subsequent RGC of Scots pine declined as root zone temperatures were lowered from ‐6°C to ‐11°C and from ‐11°C to ‐16°C. Almost no root growth was observed after exposure to ‐20°C. Shoot growth was also negatively affected by low root temperatures but less than root growth. Low root temperatures did not affect Norway spruce as much as Scots pine, although root and shoot growth of Norway spruce were reduced after exposure to the lowest test temperatures (‐16°C and ‐20°C). The length of exposure, ranging between 1 and 8 hours had no effect on subsequent growth.     

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.     

Effects of roe deer browsing and site preparation on performance of planted broadleaved and conifer seedlings when using temporary fences

Abstract

The effects of fencing and site preparation on performance of birch (Betula pendula Roth), oak (Quercus robur L.), pine (Pinus sylvestris L.) and spruce [Picea abies (L.) Karst.] seedlings were studied over four growing seasons in southern Sweden. The experiment was composed of four browsing treatments, from which roe deer (Capreolus capreolus L.) were excluded for 0, 12, 24 or 42 months, and four soil treatments. Natural browsing was combined with artificial browsing (clipping). It was demonstrated that browsing by roe deer could be isolated from browsing by moose (Alces alces L.) by selective fencing. Except for birch, browsing had little effect on seedling survival. Browsing reduced height growth by more than 100% for oak and pine, and more than 60% of pine seedling developed multiple stems. Except for oak, site preparation increased seedling survival. Inverting site preparation, in combination with or without fertilization, produced positive growth responses in seedlings, whereas patch scarification did not. There was no interaction between browsing and soil treatments on seedling performance. It was concluded that long-term protection against browsing by roe deer is needed for oak and pine, whereas short-term fences might be used for birch, and that spruce is relatively unaffected by browsing.     

Silvicultural options to promote seedling establishment on Kalmia–Vaccinium-dominated sites

Seedling growth is often hampered on sites dominated by Kalmia angustifolia. In June 2000, a trial was established on a clear-cut site in Quebec, Canada, with a high cover of Kalmia and Vaccinium species. The objectives were to evaluate how soil scarification and fertilization at the time of planting influence early growth and establishment of black spruce [Picea mariana (Mill.) BSP] and jack pine (Pinus banksiana Lamb.) seedlings. During the first 2 years, scarification reduced Kalmia cover three-fold and doubled the distance from seedlings to the nearest Kalmia stem. Scarification did not increase soil-extractable NH₄-N concentration, and reduced soil potassium, calcium and magnesium. Scarification had no effect on seedling water stress. Seedling growth improved and foliar nutrient concentrations were generally higher in scarified plots than in unscarified control plots. No differences were observed between single- and double-pass scarification for any variables except for ground-level stem diameter of seedlings, which was greater with double-pass scarification (12.1 vs 13.1 mm). Spot fertilization increased seedling growth and foliar nitrogen concentrations. Jack pine growth was greater than black spruce growth, an effect enhanced when seedlings were fertilized.     

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.     

Effect of uninucleate Rhizoctonia on Scots pine and Norway spruce seedlings

One‐year‐old container‐grown seedlings were planted in spring on clear cut areas: the Norway spruce (Picea abies) on a moist upland site (Myrtillus‐type) and Scots pine (Pinus sylvestris) on a dryish upland site (Vaccinium‐type). While still in the nursery, half of the seedlings of each species had been inoculated during the previous summer, with a uninucleate Rhizoctonia sp., a root dieback fungus. At outplanting all the seedlings appeared healthy and had a normal apical bud, although the height of the inoculated seedlings was less than that of the uninoculated control seedlings. At the end of the first growing season after planting, the mortality of inoculated Scots pine and Norway spruce seedlings was 25 and 69%, respectively. After two growing seasons the mortality of inoculated seedlings had increased to 38% for Scots pine and 93% for Norway spruce. The mortality of control seedlings after two growing seasons in the forest was 2% for Scots pine and 13% for Norway spruce. After outplanting the annual growth of inoculated seedlings was poor compared with the growth of control seedlings. These results show that, although Rhizoctonia‐affected seedlings are alive and green in the nursery, the disease subsequently affects both their survival and growth in the forest.     

Initial reactions in sapwood of Norway spruce and Scots pine after wounding and infection by Heterobasidion parviporum and H. annosum

The xylem surface of seedlings, stem material and roots of Norway spruce (Picea abies) and Scots pine (Pinus sylvestris) were inoculated with strains of Heterobasidion annosum s. str. and H. parviporum s. str. The depth of necrosis in wounded spruce increased at a linear rate for at least seven weeks of incubation, but the rate of necrotic spread was significantly faster in infected wounds. In wounded pine the necrosis was maintained at a more superficial level for several weeks. Both spruce and pine sapwood were initially infected by hyphae of both species. In spruce, the hyphae advanced at a constant rate behind the necrotic front. On the contrary, after 1–2 weeks living H. parviporum hyphae were rare in pine rays. Heterobasidion annosum hyphae survived in pine rays, phloem and tracheids, despite a heavy accumulation of phenolics and resins and were able to penetrate into the sapwood at a linear rate although slower than infections in spruce. Histochemistry and quantitative estimates demonstrated that peroxidase activity was initially higher in spruce sapwood than in pine. Within three days of incubation, the activity in spruce sapwood disappeared concurrently with deepening necrosis. However, in pine, in both control and infected samples, there was a significant increase in peroxidase activity in the area surrounding the superficial necrosis, up to the wound surface and in the cambium and phloem around the wound. After wounding and infection, the content of soluble protein increased significantly in wood of older trees but not in seedlings. Infection resulted in an increased formation of lipophilic extractives in both spruce and pine but to a significantly greater degree in the latter, whereas the amount of hydrophilic compounds decreased in both. High‐performance liquid chromatography (HPLC) analyses of lipophilic extracts showed that inoculation of pine with the two species of Heterobasidion increased the amounts of pinosylvin, its monomethylether and several other phenolics as also resinous compounds. The results obtained may be relevant in explaining the known higher resistance of Scots pine to H. parviporum.     

Water relation patterns of bare-root and container jack pine and black spruce seedlings planted on boreal cut-over sites

Water relation patterns and subsequent growth were studied on bare-root and container jack pine (Pinus banksiana Lamb.) and black spruce (Picea mariana (Mill.) B.S.P.) seedlings during the first growing season on boreal cut-over sites.Containerized seedlings of both species had greater needle conductance compared to bare-root seedlings over a range of absolute humidity deficits. Needle conductance of containerized seedlings in both species remained high during periods of high absolute humidity deficits and increasing plant moisture stress. Bare-root seedlings of both species had a greater early season resistance to water-flow through the soil-plant-atmosphere continuum (SPAC) than container seedlings. Resistance to water flow through the SPAC decreased in bare-noot seedlings of both species as the growing season progressed, and was comparable to container seedlings 9 through 14 weeks after planting. Four weeks after field planting jack pine container seedlings had greater new root development compared to bare-root seedlings, while at the end of the summer both stock types had similar new root development. Black spruce bare-root seedlings had greater new root development compared with container seedlings throughout the growing season.     

Growth in supplementarily planted Picea abies regenerations

Abstract

An experiment was established in 1978 in two Norway spruce [Picea abies (L.) Karst.] plantations in southern Sweden to study yield after mortality in patches with and without supplementarily planted (SP) seedlings. Gaps of different sizes were created by removing the originally planted seedlings. The gaps were either left unplanted or a supplementary planting was performed with one of four species [Norway spruce, Scots pine (Pinus sylvestris L.), lodgepole pine (Pinus contorta Dougl.) or hybrid larch (Larix deciduas Mill×L. Leptolepis Gord.)] 2 (at Knäred) or 6 years (at Ullasjö) after the original plantation. In 2002, most of the SP Scots pine, lodgepole pine and hybrid larch seedlings were dead or severely damaged by roe deer and moose. Survival was high among SP Norway spruces, but they had slower growth than the originally planted spruces. Growth was lower at Ullasjö than at Knäred. In Ullasjö, growth was lower in small gaps than in large gaps. Trees in original regeneration in areas surrounding unplanted gaps were larger than trees surrounding gaps with SP seedlings, which in turn were larger than originally planted trees in plots without gaps. In conclusion, because the original plantation surrounding unplanted gaps used a large part of the open space and growth of SP seedlings was slow, supplementary planting resulted in an insignificant growth increase. However, supplementary planting may increase the timber quality of trees surrounding the gaps, although this effect remains to be quantified.     

Long-term effects of stump harvesting on soil properties and tree growth in Scots pine and Norway spruce stands

Abstract

Effects of stump harvesting on the properties of surface soil and on the density, structure and growth of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) stands were estimated in a field trial in western Finland. The experiment was established in 1977 and measured in 2010. Stems and logging residues were harvested after clear-cutting, and stumps were lifted and removed from half of the experimental area. Sixteen plots were planted with pine seedlings and 16 with spruce. The main effects of stump harvesting were improved survival of planted trees and an increase in natural regeneration. No clearly negative effects were noted in the stand development. Stump harvesting had no or minimal effects on the properties of the organic layer and those of the 0- to 10-cm mineral-soil layer. Soil properties did not differ between tree species. Pine production was higher on plots with stump removal compared to plots without soil treatment.     

From planting to 26 years of age – actual growth and estimated volume scenarios for spruces and pines

The objective of this study was to compare the survival and volume of conifer stands at 26 years of age with their status at planting. Survival, growth and damage were studied in eight clear felled stands regenerated in 1972. Five of the areas were planted with Norway spruce (Picea abies (L.) Karst.) and three with Scots pine (Pinus sylvestris L.). The plantings were examined in 1972 and 1974. In 1974, the number of living undamaged planted seedlings was low (10–15%). However, the number of undamaged seedlings was supplemented by naturally regenerated conifer and birch seedlings. The total number of undamaged seedling in 1974 was equivalent to 20–30% of the number of seedlings planted. In 1998, the main species in three stands had changed from Norway spruce to Scots pine, and in one stand from Norway spruce to birches. Actual volume in 1998 for the stands was compared to stand volume generate according to five scenarios based on recommended and actual seedling number in 1972 and 1974. The actual volume was 64% of that expected if the recommended number of trees had been planted. Naturally regenerated Scots pine and Norway spruce increased stand density in 1998. The actual volume was 37% higher than the average volume in the surrounding county. On average, 36% of the trees were damaged. More than 50% of the total damage was caused by moose (Alces alces L.). For Scots pine, moose or other browsing animals damaged 30% of the trees. The results of this study indicate that the 1998 volume was higher than expected, considering the low number of undamaged seedlings in 1974. This was mainly due to the large amount of naturally regenerated plants. In addition, the results indicate that the volume could have been higher if the initial conditions had been better. Despite the low number of undamaged seedlings in 1974, seven of the eight studied stands produced a higher volume than the average stand for the region. In practise, high numbers of seedlings should be planted on scarified areas. In most cases there will be a supply of naturally regenerated seedlings.     

Boron retranslocation in Scots pine and Norway spruce

We previously traced 10B-enriched boric acid from shoots to roots to demonstrate the translocation of boron (B) in Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) seedlings. To gain a more detailed understanding of B translocation, we sought: (1) to demonstrate B retranslocation directly, by showing that foliar-applied 10B is located in the new growth after dormancy; and (2) to assess whether shoot-applied B affects growth in the long term. We applied 10B-enriched boric acid to needles of Scots pine and Norway spruce seedlings. After a dormancy period and 9 weeks of growth, small but significant increases in the 10B isotope were found in the new stem and needles of both species. In Scots pine, the total B concentration of the new stem was also increased. Both species contained polyols, particularly pinitol and inositol. Boron-polyol complexes may provide a mechanism for mobilizing B in these species. To determine the long-term effects of applied B, seedlings were grown for two growing seasons after the application of 10B to shoots. In Norway spruce, the proportion of 10B in the root systems and current needles of the harvest year was slightly higher than in the controls, and in Scots pine root systems, marginally so. The B treatment had no effect on growth of Norway spruce seedlings. In Scots pine seedlings, the B treatment caused a 33% increase in total dry mass and significantly increased the number of side branches.     

Persistence of deltamethrin against Hylobius abietis on Norway spruce seedlings

Abstract

The pine weevil Hylobius abietis L. is major threat to forest regeneration in the Nordic countries. The persistence of the deltamethrin insecticide used against pine weevil on Norway spruce seedlings was studied after the seedlings were dipped or sprayed. Insecticide application was timed to occur either before or after frozen storage. Bioassays with the stems of Norway spruce seedlings were used to determine the effect of the insecticide against feeding by the pine weevil. The measures of the control effect were reduction in area of gnawed bark and the state of health of the pine weevils. The concentration of deltamethrin decreased rapidly in seedlings, especially after spraying treatment, which did not efficiently protect seedlings against the pine weevil 6 weeks after planting. There were no signs of degradation of deltamethrin or of an effect on seedling height after frozen storage. In bioassay, the amount of deltamethrin that efficiently prevented feeding by the pine weevil was 5.5 µg g^?1 fresh weight. After one growing season in the field, about 1.76–2.24 µg g^?1 (13–15% of the initial level) of dipped deltamethrin remained in the seedlings. In seedlings treated by spraying, 0.93–0.98 µg g^?1 (7–8% of the initial level) of the deltamethrin remained. According to bioassays, these amounts were no longer sufficient to protect seedlings from feeding by the pine weevil. Therefore, in the first summer, dipping was a significantly more efficient method of application for control of pine weevils.     

Pine weevil (Hylobius abietis) damage to cuttings and seedlings of Norway spruce

Damage caused by pine weevil (Hylobius abietus L.) to planted seedlings and cuttings of Norway spruce (Picea abies (L.) Karst.) was studied at five clearcut sites in south-eastern Sweden. The main objective was to compare the two types of stock in terms of attack frequency and mortality due to pine weevil feeding. Cuttings and seedlings with the same initial stem-base diameter (4 mm) were compared. Two sites were harvested and scarified shortly before planting, two were harvested shortly before planting, but were not scarified, and one was harvested 2 years before and scarified the autumn before planting. The total mortality 5 years after planting was highest, greater than 90%, at the new, non-scarified sites, and lowest, 23%, at the old, scarified site. More than 90% of the mortality was caused by pine weevil feeding. Attack frequency and pine weevil induced mortality were significantly higher among seedlings than among cuttings. Mortality due to pine weevil damage was 4–43% higher in seedlings than in cuttings after the fifth year. Of the cuttings and seedlings that were attacked in the first year, a significantly higher frequency of the seedlings were girdled. The higher resistance of cuttings to pine weevil damage may partly explain the more rapid growth of cuttings reported in other studies. However, the causes of their higher resistance need to be further investigated. The thicker bark and needles on the stem base of the cuttings could be important in this respect.     

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.     

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.     

Heat-shock response of Pinus and Picea seedlings

The effect of a short period of elevated temperature, or heat shock, on protein synthesis was investigated in 2-day-old seedlings of jack pine (Pinus banksiana Lamb.), loblolly pine (P. taeda L.), lodgepole pine (P. contorta Dougl.), black spruce (Picea mariana (Mill.) BSP), and white spruce (P. glauca (Moench) Voss.). In all species, heat shock led to increased [(35)S]methionine incorporation into heat-shock proteins (hsp's) with molecular masses of 83 and 72 kDA. Heat shock also induced synthesis of several low molecular weight proteins that were absent from control seedlings. The low molecular weight hsp's produced by pine seedlings had molecular masses of 27, 24.6, 20.5 and 17.5 kDa, whereas those produced by spruce seedlings had molecular masses of 27.2, 19.8, 18.4, 17.2 and 16.0 kDa. All of the low molecular weight hsp's showed isoelectric variants. Heat shock led to increased [(35)S]methionine incorporation into a group of low molecular weight hsp's that were also present in control seedlings.     

Storability measures of Norway spruce and Scots pine seedlings and assessment of post-storage vitality by measuring shoot electrolyte leakage

As indoor frozen storage is increasing in forest tree nurseries it is important to have accurate methods for assessing seedling storability in autumn and methods to determine post-storage vitality. Storability of spruce (Picea abies (L.) Karst.) and pine (Pinus sylvestris L.) seedlings can be based on determination of dry matter content (DMC) of seedling shoots or by freezing shoots at –25°C and thereafter measure electrolyte leakage (SEL_diff–25). To compare these two methods we stored 1-year-old spruce and pine seedlings at different occasions during the autumn. To test if leakage of electrolytes from shoots (SEL) could indicate deteriorated vitality, we measured SEL at the end of storage. After storage seedling viability was determined in a three-week growth test, measuring shoot and root growth capacity (RGC). Determination of freezing tolerance (SEL_diff–25) before storage had a better ability to predict the outcome of storage compared to the DMC test. Measuring SEL at the end of the frozen storage period accurately indicated seedling vitality. Seedlings with SEL of 0–5% had a high survival rate whereas SEL over 10% indicated low survival and growth capacity after storage. The SEL method has a potential to become a screening test for identifying batches of seedlings that have been damaged during storage in the nursery.     

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.