The effect of late season damage on the apical bud development of Scots pine seedlings

The effect of Lygus rugulipennis Popp. feeding and artificial damage on the apical bud of Scots pine (Pinus sylvestris L.) late in the growing season was studied on two-year-old seedlings. Lygus feeding in late July and August caused inhibition of bud formation in the subsequent year's shoots, which led to loss of apical dominance and formation of interfascicular buds. Mechanical damage caused by piercing the apical bud with a needle in July, August and October produced scars and malformations on subsequent year's shoots and buds, but did not inhibit bud formation. Damage to the apical meristem could not be detected visually on the dormant bud before shoot elongation.     

Effects of soil temperature on biomass and carbohydrate allocation in Scots pine (Pinus sylvestris) seedlings at the beginning of the growing season

We studied effects of soil temperature on shoot and root extension growth and biomass and carbohydrate allocation in Scots pine (Pinus sylvestris L.) seedlings at the beginning of the growing season. One-year-old Scots pine seedlings were grown for 9 weeks at soil temperatures of 5, 9, 13 and 17 degrees C and an air temperature of 17 degrees C. Date of bud burst, and the elongation of shoots and roots were monitored. Biomass of current and previous season roots, stem and needles was determined at 3-week intervals. Starch, sucrose, glucose, fructose, sorbitol and inositol concentrations were determined in all plant parts except new roots. The timing of both bud burst and the onset of root elongation were unaffected by soil temperature. At Week 9, height growth was reduced and root extension growth was much less at a soil temperature of 5 degrees C than at higher soil temperatures. Total seedling biomass was lowest in the 5 degrees C soil temperature treatment and highest in the 13 degrees C treatment, but there was no statistically significant difference in total biomass between seedlings grown at 13 and 17 degrees C. In response to increasing soil temperature, below-ground biomass increased markedly, resulting in a slightly higher allocation of biomass to below-ground parts. Among treatments, root length was greatest at a soil temperature of 17 degrees C. The sugar content of old roots was unaffected by soil temperature, but the sugar content of new needles increased with increasing soil temperature. The starch content of all seedling parts was lowest in seedlings grown at 17 degrees C. Otherwise, soil temperature had no effect on seedling starch content.     

Infection of Scots pine seedlings by Gremmeniella abietina during summer under different inoculum potential

The connection between natural conidia dispersal of Gremmeniella abietina (A‐type) and the disease incidence and severity in first and second year conifer seedlings of Pinus sylvestris was studied in central Finland. The seedlings where exposed to natural infection for a 3‐week period throughout the growth season, followed by a 3‐week incubation period in a growth chamber to promote symptom expression. In second year seedlings the infection periods in May–June had a similar effect as regards disease outbreak and roughly half of the successful infections occurred during these periods. However, the first year seedlings were more severely diseased after the infection period in late July/early August, although the inoculum density was lower during this time compared with the period in late June/early July. This result is consistent with earlier studies. The effect of age and growth phase of the seedlings, temperature sum and the number of conidia on the disease occurrence is discussed and compared with the results of earlier studies in which artificial inoculation was employed.     

Height variation of transplanted seedlings of Scots pine in Finnish nurseries

The height variation of Scots pine (Pinus sylvestris L.) seedling stock was studied using data from annual inventories of seedling crops. The heights of 240 batches of 2-year-old and 3-year-old transplanted seedling from the years 1973–1986 grown at three nurseries in southern Finland were examined. From each batch, 100 clusters of three successive seedlings were systematically located and measured. The mean heights of the seedling batches ranged from 7.8 to 27.0 cm, with a mean of 14.1 cm. Within batch standard deviations ranged from 1.7 to 7.8 cm, with a mean standard deviation of 3.7 cm. There was considerable variation in the mean heights of seedling batches both within a given year and between years. However, estimated variance components of seedling height showed that year, batch and cluster effect each accounted for about 20 percent of the total variance, while seedling effect explained the remaining 40 percent. The implications of results for single tree and batch grading are discussed.     

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.     

Transpiration of Scots pine in Flanders growing on soil with irregular substratum

An investigation was carried out to compare the water balance of Scots pine in Flanders growing on soils with contrasted water availability. Based on sap flow measurements transpiration of Scots pine was determined for two small plots on cover sands resting on a clayey substratum of varying depths (shallow and deep). Soil water content (SWC) was relatively low (0.12–0.21 m³ m⁻³) in the upper topsoil (0–0.75 m) in both plots. However, it was always higher in the shallow plot (by 3–27%) than in the deep plot. The difference between SWC in both plots was more pronounced in the deeper soil layers (0.75–1.5 m). Sap flow was measured in seven sample pine trees on each plot from May to October 2000 using the heat field deformation (HFD) method. Transpiration of the individual trees in the deep plot was 22% lower than in trees in the shallow plot. The difference decreased to 15% after scaling up to the stand level due to a higher density of trees growing in the deep plot. It was hypothesized that higher water uptake in the shallow plot was possibly caused by structural differences between the root systems of trees growing in plots with variable soil texture. The sapwood in shallow-plot trees was 1 cm less deep than in trees growing in the deep plot (as measured by biometric and sap flow pattern methods). Sap flow radial patterns suggested a higher involvement of sinker roots for water uptake in the deep clayey substratum plot. This was in agreement with higher activity of the inner xylem in trees on the deep plot under higher evaporative demands. However, the fraction of the inner xylem to the whole-tree water supply was nearly three-fold lower than the outer xylem, which appeared to provide water presumably from the superficial roots. The fraction of these roots, estimated according to sap flow radial patterns, was around 10% higher in trees on the shallow plot. This caused 30% higher sap flow in the stem outer xylem there. Transpiration of the pine stands was limited under high evaporative demands in both plots by the low availability of soil water. The limitation was greater in the deep plot and persisted throughout the whole growing season.     

Morphological and physiological differences in Scots pine seedlings of six seed origins

One-year-old seedlings of Scots pine (Pinus sylvestris L.) offour native seed origins (Loch Maree Islands, Glengarry/GlenMorriston, Glen Affric and Abernethy), a commercial Britishseedlot, and a seedlot from Hedesunda, in middle Sweden, werecompared at monthly intervals from October 1993 to April 1994.Seedling morphology, root condition, root frost hardiness andbud dry matter were determined at each date. There were clear morphological differences among seed origins.Seedlings raised from the commercial seedlot (A70) were largerbut had a poorer root:shoot ratio than the other seed origins.Of the native pines tested, the Loch Maree Islands origin allocateda larger proportion of its photosynthate to fine roots and needlesand smaller proportion to woody structures. Seedlings raised from the commercial British seedlot tendedto have poorer bud lignification than the other origins andalso, in autumn, higher electrolyte leakage rates from its fineroots. During winter, the Swedish origin had the lowest fineroot electrolyte leakage. Seedlings of all origins showed aprogressive increase in fine root hardiness towards mid-winterwith maximum hardiness (–7°C) in January. Dehardeningoccurred over subsequent months reaching –3°C in April.Differences among origins were evident. The Swedish seedlotdeveloped greater frost resistance than the other origins, hardeningbegan earlier in autumn and dehardening began later in spring.The commercial seedlot hardened later than the other originsbut reached a similar level of frost hardiness by January. Ofthe native pines, seedlings of the Loch Maree Islands originwere slowest to develop root hardiness.     

Below-ground hydraulic constraints during drought-induced decline in Scots pine

Key message

Below-crown hydraulic resistance, a proxy for below-ground hydraulic resistance, increased during drought in Scots pine, but larger increases were not associated to drought-induced defoliation. Accounting for variable below-ground hydraulic conductance in response to drought may be needed for accurate predictions of forest water fluxes and drought responses in xeric forests.

Context

Hydraulic deterioration is an important trigger of drought-induced tree mortality. However, the role of below-ground hydraulic constraints remains largely unknown.

Aims

We investigated the association between drought-induced defoliation and seasonal dynamics of below-crown hydraulic resistance (a proxy for below-ground hydraulic resistance), associated to variations in water supply and demand in a field population of Scots pine (Pinus sylvestris L.)

Methods

Below-crown hydraulic resistance (r_bc) of defoliated and non-defoliated pines was obtained from the relationship between maximum leaf-specific sap flow rates and maximum stem pressure difference estimated from xylem radius variations. The percent contribution of r_bc to whole-tree hydraulic resistance (%r_bc) was calculated by comparing stem water potential variations with the water potential difference between the leaves and the soil.

Results

r_bc and %r_bc increased with drought in both defoliated and non-defoliated pines. However, non-defoliated trees showed larger increases in r_bc between spring and summer. The difference between defoliation classes is unexplained by differences in root embolism, and it is possibly related to seasonal changes in other properties of the roots and the soil-root interface.

Conclusion

Our results highlight the importance of increasing below-ground hydraulic constraints during summer drought but do not clearly link drought-induced defoliation with severe below-ground hydraulic impairment in Scots pine.

     

Growth regulation of Scots pine seedlings with different fertilizer compositions and regimes

Scots pine (Pinus sylvestris L.) seedlings were grown in containers filled with peat, using two different fertilizers and three different fertilizer regimes. Seedling shoot and root growth and shoot content of nitrogen, potassium and phosphorus were followed in the nursery and after outplanting in the field. Attempts to regulate growth rate by an exponential nutrient supply were not successful, but the root/shoot ratio was influenced by the fertilization regime. Internal nitrogen concentration was stable only for seedlings with low relative growth rate, while seedlings with high nutrient supply in the nursery showed strong nutrient dilution in the shoot after planting.     

Field performance of Scots pine (Pinus sylvestris L.) seedlings planted in disc trenched or mounded sites over an extended planting season

Economic pressures have driven an ever-widening period during which foresters use machines to plant Scots pine (Pinus sylvestris L.) seedlings. In Fennoscandia, this period has recently stretched to the entire growing season. To evaluate the performance of seedlings planted during this extended period, three experiments were carried out in Central and Northern Finland over 2 years. One-year-old and current-year seedlings were planted in mounds or disc-trenched furrows when soil temperatures were >0°C. When 1-year-old seedlings grown for spring planting and overwintered outdoors were planted after mid-June, more needles browned and growth was reduced, possibly because seedlings were oversized with respect to planting density and the volume of growth media. When current-year seedlings sown in spring were planted from July to November, those planted in late September and October grew less in later years than those planted earlier, but survival was unaffected. No large differences in field performance were found with respect to whether seedlings were planted in mounds or disc-trenched furrows. In conclusion, Scots pine seedlings can be machine planted in mounds or furrows during May and early June (later in the North) and then continued from early August until late September, provided climatic conditions in late spring and early autumn are typical and similar to those experienced in Central Finland.     

Rationale for growing southern pine seedlings at low seedbed densities

Although most bare-root pine seedlings in the Southern United States are grown at seedbed densities near 300/m², the density used in other regions of the world is often less than 200/m². One rationale for growing seedlings at lower seedbed densities is based on the desire to reduce the time required for successful stand establishment. Achieving a one- to two-year advancement in stand establishment can result in an additional 15 to 30 m³/ha within 15 to 20 years. Although seedling grade studies have demonstrated similar gains in volume production at ages 10 to 30 years, the findings from these studies are not widely known. The rationale in the Southern United States for growing at higher seedbed densities appears to be based on: (1) misinformation regarding the performance of morphologically improved seedlings; (2) a desire to minimize seedling and planting costs; and (3) density recommendations that are not based on volume growth in the field.     

Free amino acids and protein in Scots pine seedlings cultivated at different nutrient availabilities

Scots pine (Pinus sylvestris L.) seedlings of a provenance from northern Sweden were cultivated hydroponically for 7 weeks in a climate chamber. The nutrient solution contained either 2.5 (low-N) or 50 (high-N) mg N l(-1) with other essential elements added in a fixed optimal proportion to the nitrogen. After 5 and 7 weeks, the seedlings were analyzed for growth, total nitrogen and other essential nutrients, protein and free amino acids. Low-N seedlings grew more slowly and had higher root/shoot ratios than high-N seedlings. With respect to total nitrogen, the effect of the lower nutrient supply was mainly on the nitrogen content of the whole plant and the allocation of nitrogen among tissues, not on tissue nitrogen concentration. This was also the case for potassium, phosphorus, calcium and magnesium. The proportions by weight among these macronutrients in the whole seedlings were similar in both nutrient regimes. The proportion and concentration of sulfur were significantly lower in low-N seedlings than in high-N seedlings, because of a lower net uptake of sulfur than of other macronutrients. The shoot, needles and stem of low-N seedlings had higher concentrations of free amino acids and lower concentrations of protein than the shoot, needles and stem of high-N seedlings. Arginine dominated the pool of free amino acids in the low-N seedlings, whereas glutamine predominated in the high-N seedlings. We conclude that Scots pine seedlings accumulated soluble nitrogen as arginine when net protein synthesis was limited by factors other than nitrogen availability. Nutritional imbalance, as revealed by growth characteristics and a suboptimal proportion and concentration of sulfur in the seedlings, probably affected synthesis of S-amino acids, resulting in the diversion of assimilated nitrogen to arginine instead of protein.     

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.     

Nitrogen availability modifies the ozone responses of Scots pine seedlings exposed in an open-field system

Three-year-old Scots pine (Pinus sylvestris L.) seedlings were exposed to either ambient or elevated (1.5-1.6 x ambient) ozone concentration ([O3]) for three growing seasons in an open-field fumigation facility where they were irrigated during the growing season with a nutrient solution providing nitrogen (N) at 70 (LN treatment), 100 (control) or 150% (HN treatment) of the optimum supply rate. Treatment effects were most evident during the third year of exposure, when the ambient [O3] + HN treatment enhanced whole-plant biomass, root/shoot dry weight ratio, needle pigment concentrations and the number of chloroplast plastoglobuli in the mesophyll cells in current-year (C) needles, whereas it reduced starch accumulation in C needles and abscission of 2-year-old (C+2) needles. In the control fertilization, 3 years of exposure to elevated [O3] decreased stem-base diameter and increased K concentration and electron density of chloroplast stroma in C needles. Plants in the HN treatment exposed for 3 years to elevated [O3] had significantly lower heights, current-year main shoot length and root/shoot dry mass ratio than control plants, and increased abscission of C+2 needles. In contrast, O3-induced changes in the ultrastructure of mesophyll cells were most evident in seedlings grown for 3 years in the LN treatment. We conclude that, in Scots pine, a relatively O3-tolerant species, chronic O3 exposure leads to cumulative growth reduction, increased needle abscission and changes in carbon allocation that are strongly influenced by plant N availability.     

Changes in nitrogen level and mycorrhizal structure of Scots pine seedlings inoculated with Thelephora terrestris

? We investigated the influence of Thelephora terrestris, an ectomycorrhizal fungus, on the concentration of nitrogen in needles of Pinus sylvestris L. seedlings and their mycorrhizal structure within four seasons of growth.

? Seedlings were grown in four treatments: (I) inoculated + 0.030 g N, (II) inoculated + 0.045 g N, (III non-inoculated + 0.030 g N, (IV) non-inoculated + 0.045 g N under laboratory conditions and later planted in post-agricultural land.

? For inoculated treatments, statistically significant differences in N concentration of needles were observed for two- and four-year-old seedlings. The number of ectomycorrhizae and concentration of N in needles were negatively correlated. After four seasons of growth the greatest numbers of ectomycorrhizae were present on seedlings from treatment II, and the smallest on seedlings from treatment IV.

? Two-year-old seedlings had been colonized by at least six different fungal taxa, i.e. Cenococcum geophilum, Hebeloma crustuliniforme, Rhizopogon sp., Suillus bovinus, Thelephora terrestris and a non-identified one. Treatments I and II were dominated by the ectomycorrhizae of T. terrestris and their number was significantly greater than in treatments III or TV. On four-year-old seedlings ectomycorrhizae of T. terrestris were dominated in all treatments, while ectomycorrhizae of Hebeloma sp. were not found.

     

Regulation of organic and inorganic nitrogen uptake in Scots pine (Pinus sylvestris) seedlings

Plants possess regulatory mechanisms that enhance nitrogen (N) uptake under conditions of spatial and temporal variation in N availability. Study of regulatory mechanisms has focused almost exclusively on the uptake of inorganic N sources (i.e., ammonium (NH4+), nitrate (NO3-). Several lines of evidence, however, suggest that amino acids may constitute a potential source of N for a number of plant species, including conifers. In the present study, we investigated the uptake of amino acids and inorganic N in Scots pine (Pinus sylvestris L.) seedlings grown at different N concentrations. We compared the uptake rate of the individual N sources using U-[13C2], [15N]-glycine, U-[13C6], [15N4]-arginine, 15NH4, or 15NO3, and tested the short-term effect of N supply on the uptake rate of glycine, arginine and in field-grown Scots pine seedlings. Our data indicate that Scots pine seedlings can absorb substantial amounts of N in the form of intact arginine and glycine molecules. The data also suggest that Scots pine seedlings down-regulate their uptake of NH4+-N and arginine-N, but not of glycine-N in response to increased endogenous N concentrations.     

Survival of Scots pine seedlings after biological and chemical control of damping-off in plastic greenhouses

Survival of pine seedlings was tested in greenhouses where substrates were infected with Rbizoctonia, Fusarium or Rbizoctonia with Fusarium. To reduce the pathogen menace biological (sterile sawdust, Tricboderma or sawdust with Tricboderma) or chemical (seed protection) treatments were performed. Incorporation of sawdust or sawdust with Tricboderma 20 days before sowing increased the survival of seedlings and it was comparable to protection with Nemispor for Fusarium-menace, with Funaben T for Rbizoctonia menace and Oxafun T or Karbochin for both pathogens.     

Lignin deposition during earlywood and latewood formation in Scots pine stems

Lignin deposition at consecutive secondary wall thickening stages of early and late xylem cells during annual ring wood formation in Scots pine (Pinus sylvestris L.) stems was studied. Lignin patterns, isolated by thioglycolic acid method, consisted of alcohol-soluble (LTGA-I) and alkali-soluble (LTGA-II) fractions. The sum of two fractions, being the total lignin content, gradually increased in the course of lignification. However, the increments of lignin amount at each development stage of early and late tracheids were different. The intensity of lignin deposition increased in the course of earlywood tracheid maturation and decreased toward the end of latewood cell differentiation. The deposition of two lignin fractions in each layer of forming wood also occurred oppositely. The increment of LTGA-I descended, whereas that of LTGA-II increased from the beginning to the end of early xylem lignification. In contrast, LTGA-I increment dropped, whereas LTGA-II rose during late xylem lignification. Gel permeation chromatography showed that the lignins, formed at the beginning of lignification, were more homogeneous and had higher molecular weight compared with the lignins at the end of cell differentiation. Besides, the content of cellulose, estimated as the residue after lignin isolation, and of cell wall substances, presented as cell wall cross-section areas, at consecutive maturation stages of early and late xylem cells have been found to be different. The data show that lignin deposition occurred in different conditions and with opposite dynamics during early and late xylem formation.