Seedling shoot,needle and bud development in three provenances of Pinus sylvestris and Pinus contorta cultivated in northern Sweden

The patterns of current‐year shoot, needle and terminal bud elongation in seedlings of three Scots pine (Pinus sylvestris L.) and three lodgepole pine (Pinus contorta Dougl. var. latifolia Engelm.) provenances were compared during the third and fourth growing seasons after planting. Lodgepole pine produced longer shoots and buds than did Scots pine, mainly because lodgepole pine formed more stem units and elongated at a faster rate. Stem unit length and the duration of shoot and bud elongation differed relatively little between species and provenances. Lammas or polycyclic growth occurred in some lodgepole pine provenances, but not in any Scots pine provenance, and was associated with enhanced shoot elongation. Needle elongation commenced earlier, proceeded at a faster rate, and was greater in lodgepole pine than in Scots pine, but ceased about the same time in all species and provenances. The heat sum required to attain 50% of final length was lower for shoots and needles in lodgepole pine than in Scots pine, and for shoots in northern provenances than in southern ones. Mitotic activity in the apical meristem of the terminal bud, which occurred less than one week after the seedlings were free from snow, started and ceased about the same time in each species, but was higher in lodgepole pine than in Scots pine early in the shoot elongation period.     

Seasonal differences in freezing stress resistance of needles of Pinus nigra and Pinus resinosa: evaluation of the electrolyte leakage method

Seasonal changes in freezing stress resistance of needles of red pine (Pinus resinosa Ait.) and Austrian pine (Pinus nigra Arnold) trees were measured by an electrolyte leakage method and by visual observation. During most of the year, freezing stress resistance determined by the two methods gave similar results. The electrolyte leakage method provided a good estimate of seasonal changes in freezing stress resistance except for red pine needles in their most winter-hardy state. To obtain a reliable estimate of freezing stress resistance in winter-hardy red pine needles it was necessary to combine the electrolyte leakage method with visual observations. When red pine needles survived exposure to -80 degrees C or lower, electrolyte leakage was never more than 30% even when the needles were exposed to a slow freeze-thaw stress of -196 degrees C. However, rapid freezing of red pine needles to -196 degrees C resulted in electrolyte leakage of over 80%. Red pine needles attained a much higher freezing stress resistance during the winter than Austrian pine. Red pine needles also acclimated and deacclimated faster than Austrian pine needles. An index of injury was developed based on the electrolyte leakage method ((R(2) + R(1))/2, where R(1) is the minimum % electrolyte leakage from noninjured tissue and R(2) is the maximum % electrolyte leakage at the highest injury) that reliably predicted freezing stress resistance of pine needles for most of the year. Important aspects for developing a successful index of injury for pine needles are: use of cut needles, vacuum infiltration and shaking during incubation in water.We conclude that: (1) during cold acclimation the cell wall properties of the pine needles changed and these changes, which appeared to differ in the two species, might explain the very low leakage of electrolytes from winter-hardy needles of red pine; (2) pine needles survive winter by developing the ability to tolerate extracellular ice formation, because after rapid freezing the needles were severely injured; and (3) red pine is adapted to a shorter growing season and colder winters than Austrian pine.     

Genetic and environmental control of seasonal carbohydrate dynamics in trees of diverse Pinus sylvestris populations

We explored environmental and genetic factors affecting seasonal dynamics of starch and soluble nonstructural carbohydrates in needle and twig cohorts and roots of Scots pine (Pinus sylvestris L.) trees of six populations originating between 49 degrees and 60 degrees N, and grown under common garden conditions in western Poland. Trees of each population were sampled once or twice per month over a 3-year period from age 15 to 17 years. Based on similarity in starch concentration patterns in needles, two distinct groups of populations were identified; one comprised northern populations from Sweden and Russia (59-60 degrees N), and another comprised central European populations from Latvia, Poland, Germany and France (49-56 degrees N). Needle starch concentrations of northern populations started to decline in late spring and reached minimum values earlier than those of central populations. For all populations, starch accumulation in spring started when minimum air temperature permanently exceeded 0 degrees C. Starch accumulation peaked before bud break and was highest in 1-year-old needles, averaging 9-13% of dry mass. Soluble carbohydrate concentrations were lowest in spring and summer and highest in autumn and winter. There were no differences among populations in seasonal pattern of soluble carbohydrate concentrations. Averaged across all populations, needle soluble carbohydrate concentrations increased from about 4% of needle dry mass in developing current-year needles, to about 9% in 1- and 2-year-old needles. Root carbohydrate concentration exhibited a bimodal pattern with peaks in spring and autumn. Northern populations had higher concentrations of fine-root starch in spring and autumn than central populations. Late-summer carbohydrate accumulation in roots started only after depletion of starch in needles and woody shoots. We conclude that Scots pine carbohydrate dynamics depend partially on inherited properties that are probably related to phenology of root and shoot growth.     

Temporal variation in nutrient concentrations of Pinus sylvestris needles

Temporal variation in nutrient concentrations of Scots pine (Pinus sylvestris L.) needles was studied during a three‐year period in three stands of differing stages of development. Concentrations of N, P and K varied significantly between years; this variation was related to differences in needle dry weight. Concentrations of all measured nutrients (N, P, K, Ca, Mg, Mn, Cu, Zn, Fe, B) and Al varied between seasons; this variation was related to nutrient mobility and the annual physiological cycle. Concentrations of the mobile nutrients N, P and K decreased in spring and early summer during shoot and needle elongation and increased in late summer and autumn during needle senescence and litterfall. Concentrations of Mg, Cu, Zn and B followed somewhat similar patterns. The poorly mobile nutrients Ca, Mn and Fe accumulated gradually in needles during each growing season. Needle nutrient concentrations were stable during the nonactive period.     

Seasonal acclimation of photosystem II in Pinus sylvestris. II. Using the rate constants of sustained thermal energy dissipation and photochemistry to study the effect of the light environment

Photosynthesis in evergreen conifers is characterized by down-regulation in autumn and rapid up-regulation in spring. This seasonal pattern is largely driven by temperature, but the light environment also plays a role. In overwintering Scots pine (Pinus sylvestris L.) trees, PSII is less down-regulated and recovers faster from winter stress in shaded needles than in needles exposed to full sunlight. Because the effect of light on the seasonal acclimation of PSII has not been quantitatively studied under field conditions, we used the rate constants for sustained thermal energy dissipation and photochemistry to investigate the dynamics and kinetics of the seasonal acclimation of PSII in needles exposed to different light environments. We monitored chlorophyll fluorescence and needle pigment concentration during the winter and spring in Scots pine seedlings growing in the field in different shading treatments, and within the crowns of mature trees. The results indicated that differences in acclimation of PSII in overwintering Scots pine among needles exposed to different light environments can be chiefly attributed to sustained thermal dissipation. We also present field evidence that zeaxanthin-facilitated thermal dissipation and aggregation of thylakoid membrane proteins are key mechanisms in the regulation of sustained thermal dissipation in Scots pine trees in the field.     

Structure and abundance of arachnid communities in Scots and lodgepole pine plantations

Between April and September 1994, six plots within Aberfoyle Forest in Scotland, three of Scots (Pinus sylvestris) and three of lodgepole pine (Pinus contorta), were surveyed for spiders and harvestmen, as these are predators of several forest invertebrate pests.A total of 64 species of spider were trapped in Scots and lodgepole pine; 56 in Scots pine and 32 in Lodgepole pine. The most abundant spiders, both in Scots and Lodgepole pine, were Linyphiidae.Scots pine plots were floristically more diverse and structurally more complex than lodgepole pine plots. More species of spiders and harvestmen (midsummer only) were found at ground level in Scots pine than lodgepole pine plots. More species and a greater numbers of spiders were found in the canopy of Scots pine than in that of lodgepole pine. However, no differences in the abundance of harvestmen were found between the canopies of Scots and lodgepole pine. Spider species diversity was greatest in Scots pine.     

Impacts of seasonal air and soil temperatures on photosynthesis in Scots pine trees

Seasonal courses of light-saturated rate of net photosynthesis (A360) and stomatal conductance (gs) were examined in detached 1-year-old needles of Scots pine (Pinus sylvestris L.) from early April to mid-November. To evaluate the effects of soil frost and low soil temperatures on gas exchange, the extent and duration of soil frost, as well as the onset of soil warming, were manipulated in the field. During spring, early summer and autumn, the patterns of A360 and gs in needles from the control and warm-soil plots were generally strongly related to daily mean air temperatures and the frequency of severe frost. The warm-soil treatment had little effect on gas exchange, although mean soil temperature in the warm-soil plot was 3.8 degrees C higher than in the control plot during spring and summer, indicating that A360 and gs in needles from control trees were not limited by low soil temperature alone. In contrast, prolonged exposure to soil temperatures slightly above 0 degrees C severely restricted recovery of A360 and especially gs in needles from the cold-soil treatment during spring and early summer; however, full recovery of both A360 and gs occurred in late summer. We conclude that inhibition of A360 by low soil temperatures is related to both stomatal closure and effects on the biochemistry of photosynthesis, the relative importance of which appeared to vary during spring and early summer. During the autumn, soil temperatures as low as 8 degrees C did not affect either A360 or gs.     

Nitrate metabolism in Scots pine seedlings during their first growing season

Fluctuations in nitrate reductase activity (NRA), nitrate, nitrite, protein and total nitrogen content of bare-root Scots pine seedlings (Pinus sylvestris L.) raised outdoors were investigated during the first growing season. Nitrate reductase activity was higher in roots than in needles, whereas NO(3) (-) content was higher in needles than in roots and increased in both from June to October. Nitrate reductase activity in roots correlated more closely with NO(3) (-) N in the soil than did NO(3) (-) in the roots. In autumn, there was a closer correlation between foliar NRA and NO(3) (-) in the needles than with NO(3) (-)-N in the soil. Nitrite was not detected in the seedlings during the growing season. Total nitrogen content decreased toward the autumn, whereas protein content initially decreased but increased again in autumn. Acrylic netting placed above the seedlings increased both air and soil temperatures and apparently accelerated the use of nitrate.     

Tap root formation and early root/shoot ratio of Pinus Contorts and Pinus sylvestris

Scots pine and different provenances of lodgepole pine were compared in regard to root characteristics and the distribution of biomass within individual trees. Tap root formation was more frequent and the root/shoot ratio was higher on Scots pine than on lodgepole pine but significant variation of these traits was found between provenances of planted lodgepole pine. A non‐destructive method for estimation of root biomass was developed.     

Frost damage in Pinus sylvestris L. stems assessed by chlorophyll fluorescence in cortical bark chlorenchyma

? Several techniques were used to investigate the threshold minimum temperature inducing severe frost damage during winter in Scots pine (Pinus sylvestris L.) needles and stems from southern provenances in the Iberian Peninsula.

? Chlorophyll (Chl) fluorescence, electrolyte leakage (EL), visual scoring (VS), and the normalized difference vegetation index (NDVI) were compared.

? Chl fluorescence, and in particular the maximum potential photosystem II (PSII) efficiency — estimated through the dark-acclimated F_V/F_M ratio-, was found to be a simple, non-destructive indicator of freezing tolerance in needles, as expected.

? Moreover, the existence of a Chl-containing tissue, the cortical bark chlorenchyma, allowed us the use of Chl fluorescence as a new non-destructive indicator of frost damage in live tissues of Pinus sylvestris L. stems.

? Freezing tolerance values of southern provenances are compared with those found by other authors in the northern distribution limit of the species.

     

Nutrient retranslocation within the foliage of Pinus sylvestris

During the period 1983-1987, retranslocation of N, P, K, Ca, Mg, Mn, Zn, Fe, B and Al within the foliage of Scots pine (Pinus sylvestris L.) was studied in sapling, pole-stage and mature stands in eastern Finland. Needle concentrations of the mobile nutrients N, P, and K varied seasonally because of retranslocation. In unfertilized plots, needle contents of N, P and K decreased 62-92% during senescence and needle dry weight decreased 19-51%. In some years, needle contents of Ca, Mn, Zn, Fe, B and Al increased during senescence. Retranslocation was more efficient from needles with greater dry weight. In the fertilized, pole-stage stand, needle dry weight did not decrease at senescence four years after fertilization and nutrient content decreased less than in the other years. In the mature stand, there were no clear differences in needle dry weight and nutrient changes during senescence between years or between unfertilized and fertilized plots.     

The effects of nitrogen fertilization on the ultrastructure of mesophyll cells of conifer needles in northern Finland

Cytological injuries in needles of Scots pine (Pinus sylvestris L.) and Norway spruce [Picea abies (L.) Karst.] were studied at light and electron microscope level in northern Finland. Needle samples were taken from fertilized (mainly with nitrogen) and nonfertilized trees during winter 1980. The results are discussed in relation to the severe climate and growth response of the trees.     

Responses of Pinus massoniana and Pinus taeda to freezing in temperate forests in central China

Abstract

Masson pine (Pinus massoniana Lamb.), a native species widely distributed in temperate forests in central China, and Loblolly pine (Pinus taeda L.), an exotic tree species introduced to China from southeastern United States, are dominant evergreen conifers that play a pivotal role in maintaining forest structure and functions for the region. We examined the effects of freezing on these species with chlorophyll fluorescence and electrolyte leakage using both field- and laboratory-based experiments in September 2009 and January 2010, respectively. We found that freezing could cause a greater impact on the Loblolly pine than the Masson pine. Although the two species showed similar values of F _v /F _m and electrolyte leakage before freezing, the Masson pine needles showed lower F _v /F _m and higher electrolyte leakage ratios than those of the Loblolly pine when treated in low temperatures (?15 to 0°C). We also found that cold-acclimation was crucial for both species to adapt to low temperatures with the F _v /F _m ratio decreased approximately by 80% in the first freezing hour for the non-acclimated needles of both species while the cold-acclimated needles showed little changes in the F _v /F _m ratio. This finding is also supported by our measurements of electrolyte leakage. These results suggest that the Loblolly pine could be more susceptible to freezing damages than the Masson pine in central China.     

Relationship between temperature, respiratory loss of sugar and premature dehardening in dormant Scots pine seedlings

Increased intracellular sugar concentration is an important contributor to the increased cold tolerance of conifers in winter. This study examines the extent to which wintertime respiratory loss of sugars leads to premature dehardening. Two-year-old seedlings of Scots pine (Pinus sylvestris L.), grown and cold-hardened in the field, were exposed to different temperature regimes for 16 weeks while dormant. To minimize short-term carry-over effects, after the temperature treatments, all seedlings were conditioned to 5.5 degrees C and watered before the assessment of non-structural carbohydrates and cold tolerance. Needle sugar concentration was decreased by 54, 32, 21 and 9% following treatment at 5.5, 0, -1.5 and -8.5 degrees C, respectively. Sugar concentration did not decrease as much in root tissues as in needles because starch was mobilized in roots. Cold tolerance of needles was analyzed by controlled freezing, and the temperature causing an initial 10% damage (LT(10)) was plotted as a function of needle sugar concentration, revealing a strong, linear relationship. When one-third of the initial sugars had been consumed, LT(10) had increased from -24.5 to -16.5 degrees C, and when one half had been consumed, LT(10) had increased to -10 degrees C. Consequences of these findings for the field performance of conifers are discussed in relation to climatic variation and change.     

Effects of low night temperature and light on chlorophyll fluorescence of field-grown seedlings of Scots pine (Pinus sylvestris L.)

In vivo chlorophyll fluorescence kinetics was observed in naturally regenerated and planted Scots pine (Pinus sylvestris L.) seedlings growing at two adjacent sites in northern Sweden. Some seedlings were shaded from direct solar radiation and some were protected from frost at night. Air temperature and solar radiation were monitored. In vivo chlorophyll fluorescence kinetics of current-year needles was measured at both room temperature and 77 K. On clear days during August, variable fluorescence of photosystem II of needles from exposed, naturally regenerated seedlings was reduced by night frosts, whereas frost caused little change in fluorescence of needles from shaded seedlings. Overnight frost-protection reduced the inhibition of photosystem II by direct sunlight. Recovery from photoinhibitory damage occurred on cloudy days. In September, the decline of variable fluorescence of photosystem II was more pronounced in unshaded than in shaded seedlings, and coincided with frosts at night. In addition, the reduction in variable fluorescence was larger for planted seedlings than for naturally regenerated seedlings. It is suggested that the inhibition of photosystem II activity was caused by an interaction between low temperatures and light. Natural night frosts are proposed to increase the susceptibility to photoinhibition in the following day(s).     

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.     

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.     

Changes in nutrient concentrations and nutrient release in decomposing needle litter in monocultural systems of Pinus contorta and Pinus sylvestris—a comparison and synthesis

Scots pine (Pinus sylvestris) and lodgepole pine (Pinus contorta) needle litters were compared in terms of nutrient composition and its change during decomposition. Initial nutrient composition differed between the species, with lodgepole pine needle litter having significantly higher concentrations of P, Mg and Mn. However, no difference was found for concentrations of N, Ca or K. Increases in concentrations of N, P and K during decomposition were significant in both litter types. For Ca the pattern of concentration changes followed a quadratic function as decomposition proceeded. Concentrations of Mg and Mn decreased in lodgepole pine needle litter. In Scots pine litter there was also an initial decrease, but it was followed by an increase in most incubations. For both Mg and Mn, changes in concentrations during decomposition differed significantly between species. In the late decomposition stages, concentrations of Mg and Mn became similar in both litter types. Nutrient concentrations generated by the models were compared with those of the humus (F and H) layer in the stands. The model was quite accurate in predicting concentrations of N and P for both species and the concentration of Mg for lodgepole pine. By contrast, it was not accurate in predicting concentrations of Ca and Mn. Nutrient release was estimated for the two species using both measured litterfall data and long‐term estimates, and regression models were used to predict concentration changes. Rates of release of P, Mg and Mn in the lodgepole pine stands were found to be about twice as high compared with those in Scots pine. Calcium was also released to a greater extent although the difference was not significant.     

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.