Responses of leaf structure and photosynthetic properties to intra-canopy light gradients: a common garden test with four broadleaf deciduous angiosperm and seven evergreen conifer tree species.

Spectra of leaf traits in northern temperate forest canopies reflect major differences in leaf longevity between evergreen conifers and deciduous broadleaf angiosperms, as well as plastic modifications caused by within-crown shading. We investigated (1) whether long-lived conifer leaves exhibit similar intra-canopy plasticity as short-lived broadleaves, and (2) whether global interspecific relationships between photosynthesis, nitrogen, and leaf structure identified for sun leaves adequately describe leaves differentiated in response to light gradients. We studied structural and photosynthetic properties of intra-tree sun and shade foliage in adult trees of seven conifer and four broadleaf angiosperm species in a common garden in Poland. Shade leaves exhibited lower leaf mass-per-area (LMA) than sun leaves; however, the relative difference was smaller in conifers than in broadleaves. In broadleaves, LMA was correlated with lamina thickness and tissue density, while in conifers, it was correlated with thickness but not density. In broadleaves, but not in conifers, reduction of lamina thickness was correlated with a thinner palisade layer. The more conservative adjustment of conifer leaves could result from a combination of phylogenetic constraints, contrasting leaf anatomies and shoot geometries, but also from functional requirements of long-lived foliage. Mass-based nitrogen concentration (N(mass)) was similar between sun and shade leaves, and was lower in conifers than in deciduous broadleaved species. Given this, the smaller LMA in shade corresponded with a lower area-based N concentration (N(area)). In evergreen conifers, LMA and N(area) were less powerful predictors of area-based photosynthetic rate (A (max(area))) in comparison with deciduous broadleaved angiosperms. Multiple regression for sun and shade leaves showed that, in each group, A (max(mass)) was related to N(mass) but not to LMA, whereas LMA became a significant codeterminant of A (max(mass)) in analysis combining both groups. Thus, a fundamental mass-based relationship between photosynthesis, nitrogen, and leaf structure reported previously also exists in a dataset combining within-crown and across-functional type variation.

Acclimation of respiratory temperature responses in northern and southern populations of Pinus banksiana.

Temperature acclimation of respiration may contribute to climatic adaptation and thus differ among populations from contrasting climates. Short-term temperature responses of foliar dark respiration were measured in 33-yr-old trees of jack pine (Pinus banksiana) in eight populations of wide-ranging origin (44-55 degrees N) grown in a common garden at 46.7 degrees N. It was tested whether seasonal adjustments in respiration and population differences in this regard resulted from changes in base respiration rate at 5 degrees C (R(5)) or Q(10) (temperature sensitivity) and covaried with nitrogen and soluble sugars. In all populations, acclimation was manifest primarily through shifts in R(5) rather than altered Q(10). R(5) was higher in cooler periods in late autumn and winter and lower in spring and summer, inversely tracking variation in ambient air temperature. Overall, R(5) covaried with sugars and not with nitrogen. Although acclimation was comparable among all populations, the observed seasonal ranges in R(5) and Q(10) were greater in populations originating from warmer than from colder sites. Population differences in respiratory traits appeared associated with autumnal cold hardening. Common patterns of respiratory temperature acclimation among biogeographically diverse populations provide a basis for predicting respiratory carbon fluxes in a wide-ranging species.

Climate warming will reduce growth and survival of Scots pine except in the far north.

Tree growth and survival were assessed in 283 populations of Scots pine (Pinus sylvestris L.) originating from a broad geographic range and grown at 90 common-garden experimental sites across Europe, and in 101 populations grown at 14 sites in North America. Growth and survival were analysed in response to climatic transfer distance, the difference in mean annual temperature (MAT) between the site and the population origin. Differences among populations at each site, and across sites for regional groups of populations, were related to climate transfer distance, but in opposite ways in the northern vs. southern parts of the species range. Climate transfers equivalent to warming by 1-4 degrees C markedly increased the survival of populations in northern Europe (>or= 62 degrees N, < 2 degrees C MAT) and modestly increased height growth >or= 57 degrees N but decreased survival at < 62 degrees N and modestly decreased height growth at < 54 degrees N latitude in Europe. Thus, even modest climate warming will likely influence Scots pine survival and growth, but in distinct ways in different parts of the species range.

Whole-plant allocation to storage and defense in juveniles of related evergreen and deciduous shrub species.

In evergreen plants, old leaves may contribute photosynthate to initiation of shoot growth in the spring. They might also function as storage sites for carbohydrates and nitrogen (N). We hence hypothesized that whole-plant allocation of carbohydrates and N to storage in stems and roots may be lower in evergreen than in deciduous species. We selected three species pairs consisting of an evergreen and a related deciduous species: Mahonia aquifolium (Pursh) Nutt. and Berberis vulgaris L. (Berberidaceae), Prunus laurocerasus L. and Prunus serotina Ehrh. (Rosaceae), and Viburnum rhytidophyllum Hemsl. and Viburnum lantana L. (Adoxaceae). Seedlings were grown outdoors in pots and harvested on two dates during the growing season for the determination of biomass, carbohydrate and N allocation ratios. Plant size-adjusted pools of nonstructural carbohydrates in stems and roots were lower in the evergreen species of Berberidaceae and Adoxaceae, and the slope of the carbohydrate pool vs plant biomass relationship was lower in the evergreen species of Rosaceae compared with the respective deciduous species, consistent with the leading hypothesis. Pools of N in stems and roots, however, did not vary with leaf habit. In all species, foliage contained more than half of the plant's nonstructural carbohydrate pool and, in late summer, also more than half of the plant's N pool, suggesting that in juvenile individuals of evergreen species, leaves may be a major storage site. Additionally, we hypothesized that concentration of defensive phenolic compounds in leaves should be higher in evergreen than in deciduous species, because the lower carbohydrate pool in stems and roots of the former restricts their capacity for regrowth following herbivory and also because of the need to protect their longer-living foliage. Our results did not support this hypothesis, suggesting that evergreen plants may rely predominantly on structural defenses. In summary, our study indicates that leaf habit has consequences for storage economics at the whole-plant level, with evergreen shrub species storing less carbohydrates (but not N) per unit plant biomass than deciduous species.

Temperature and ontogeny mediate growth response to elevated CO2 in seedlings of five boreal tree species.

We tested the extent to which growth responses to elevated carbon dioxide (CO2 ) are temperature-dependent and change through early seedling ontogeny among boreal tree species of contrasting relative growth rates (rgr). Populus tremuloides Michx, Betula papyrifera Marsh, Larix laricina (Du Roi) K. Koch, Pinus banksiana Lamb., and Picea mariana (Mill.) B.S.P. were grown from seeds for 3 months in controlled-environment chambers at two CO2 concentrations (370 and 580 µmol mol-1 ) and five temperature regimes of 18/12, 21/15, 24/18, 27/21 and 30/24°C (light/dark). Growth increases in response to CO2 enrichment were minimal at the lowest temperature and maximal at 21/15°C for the three conifers and at 24/18°C or higher for the two broadleaved species, corresponding with differences in optimal temperatures for growth. In both CO2 treatments, rgr among species and temperatures correlated positively with leaf area ratio (lar) (r⩾0·90, P<0·0001). However, at a given lar, rgr was higher in elevated CO2 , owing to enhanced whole-plant net assimilation rate. On average in all species and temperatures at a common plant mass, CO2 enrichment increased rgr (9%) through higher whole-plant net assimilation rate (22%), despite declines in lar in high CO2 (11%). Reductions in lar are thus an important feedback mechanism reducing positive plant growth responses to CO2 . Proportional allocation of dry mass to roots did not vary between CO2 treatments. Early in the experiment, proportional increases in plant dry mass in elevated CO2 were larger in faster-growing Populus tremuloides and B. papyrifera than in the slower-growing conifers. However, growth increases in response to CO2 enrichment fell with time for broadleaved species and increased for the conifers. With increasing plant size over time, compensatory adjustments to CO2 enrichment in the factors that determine rgr, such as lar, were much larger in broadleaves than in conifers. Thus, the hypothesis that faster-growing species are more responsive to elevated CO2 was not supported, given contrasting patterns of growth response to CO2 with increasing plant size and age.

Light environment alters response to ozone stress in seedlings of Acer saccharum Marsh, and hybrid Populus L.: I. In situ net photosynthesis, dark respiration and growth.

Hybrid poplar (Populus tristis Fisch. ×P. balsamifera L., cv. Tristis) and sugar maple (Acer saccharum Marsh.) seedlings were grown under contrasting light and ozone treatments to investigate the role of the light environment in their response to chronic ozone stress. In consecutive growth chamber experiments, cuttings of shade-intolerant poplar and 3-yr-old seedlings of shade-tolerant sugar maple were grown in pots for 6 and 10 wk, respectively, under shaded, low light irradiance (c. 2.5 mol m-2 d-1 PPFD or 7% of full sunlight) and six-fold greater irradiance (c. 16.6 mol m-2 d-1 PPFD or 45% of full sunlight) in combination with low (< 10 nl 1-1 ) and elevated levels of ozone (c. 99-115 nl 1-1 ). In unshaded poplar plants, ozone exposure reduced root dry mass by 33% at final harvest, while shaded plants had no such response. By comparison, sugar maple root dry mass was reduced by ozone in shaded plants by 10%, but was unaffected by ozone in unshaded plants. In poplar, leaf area: plant dry mass ratios were unaffected by ozone, whereas in sugar maple ozone-exposed plants had a 24% lower leaf area: plant dry mass ratio in the shaded treatment. In shade-grown sugar maple, ozone doubled dark respiration rates of leaves, but in unshaded seedlings ozone had no effect on respiration. In comparison, in poplar plants ozone exposure resulted in greater increases in dark respiration under unshaded than shaded conditions. In unshaded plants, ozone treatment resulted in lower in situ net photosynthesis in poplar, but not in sugar maple. Overall, shade-grown sugar maple appeared more sensitive to ozone stress than unshaded plants in terms of lower leaf area: plant dry mass ratio and root growth and higher leaf respiration. In poplar on the other hand, root growth, leaf respiration and photosynthesis were more affected by ozone in unshaded than in shaded plants. These findings suggest that shade-grown sugar maple and unshaded poplar may experience greater reductions in carbon gain and growth under elevated levels of ozone than plants under the opposite light conditions.

Light environment alters response to ozone stress in seedlings of Acer saccharum Marsh, and hybrid Populus L.: II. Diagnostic gas exchange and leaf chemistry.

Diagnostic gas exchange measurements and foliar chemical assays were conducted on hybrid poplar (Populus tristis Fisch. ×P. balsamifera L. cv. Tristis) and sugar maple (Acer saccharum Marsh.) seedlings grown under contrasting light and ozone treatments. Seedlings were grown in low irradiance (c. 2.5 mol m-2 d-1 ) and six-fold greater irradiance (c. 16.6 mol m-2 d-1 ) in combination with low (< 10 nl I -l ) and elevated (99-115 nl 1-1 ) ozone. Analysis of light response curves showed ozone-induced reductions in photosynthetic capacity and quantum yield for unshaded poplar and shaded sugar maple, but not the contrasting light treatments. Photosynthesis at saturating CO2 concentrations was decreased in the elevated ozone treatment in both the unshaded and shaded poplar and in shaded sugar maple. Poplar had significant reductions in chlorophyll concentration due to ozone exposure in both unshaded and shaded treatments. Older leaves of unshaded poplar plants had significantly greater reductions in chlorophyll levels due to ozone than older leaves of shaded plants. In maple, only shade-grown leaves had significant decreases in chlorophyll concentration due to ozone exposure. The diagnostic gas exchange measurements in conjunction with chlorophyll measurements indicate that in hybrid poplar, unshaded leaves may be more sensitive to ozone than shade leaves, while in sugar maple, shade leaves are more sensitive to ozone. For hybrid poplar a decrease in photosynthetic capacity, quantum yield and chlorophyll concentration in the unshaded, moderately high light environment due to elevated ozone is consistent with prior studies. The results indicating that sugar maple seedlings may be more detrimentally affected by elevated ozone in the lower light environment may have serious implications for this and other shade-adapted species with respect to their performance in an understorey environment.

Light environment alters response to ozone stress in seedlings of Acer saccharum Marsh, and hybrid Populus L.: III. Consequences for performance of gypsy moth.

Both light conditions and ozone fumigation alter the chemical composition of tree foliage and are thus likely to influence tree-insect interactions. We investigated the direct and interactive effects of light environment and ozone exposure on the performance of gypsy moth (Lymantria dispar L.) larvae reared on hybrid poplar (Populus tristis Fisch. ×P. balsamifera L. cv. Tristis) and sugar maple (Acer saccharum Marsh). We used a split-plot experimental design (light nested within ozone) and fourth-instar bioassays to calculate standard indices of insect growth and feeding performance. For insects fed poplar, consumption, growth and processing efficiencies were affected more by light environment than by ozone. Larvae ate and grew less on high-light foliage, responses attributable to higher levels of phenolic glycosides in those leaves. For insects fed maple, no significant effects of light, ozone, or light x ozone were observed. These results demonstrate that light environment and ozone pollution can alter the dynamics of interactions between trees and associated insects and that responses are species-specific.

Primary and secondary host plants differ in leaf-level photosynthetic response to herbivory: evidence from Alnus and Betula grazed by the alder beetle, Agelastica alni.

Field-grown trees of Alnus incana (L.) Moench, Alnus glutinosa (L.) Geartner and Betula pendula Roth displayed pronounced differences in responses of light-saturated net photosynthesis (Asat ) to herbivory by the alder beetle (Agelastica alni L., Galerucinae), a specialized insect which primarily defoliates alders. We found that photosynthetic rates of grazed leaves increased following herbivory in Alnus but not in Betula. Area- and mass-based Asat of grazed leaves declined linearly with increasing amount of leaf perforation in B. pendula, by as much as 57%. By contrast Alnus glutinosa and Alnus incana increased area-based rates of Asat by 10-50% at all levels of leaf grazing. Given increased Asat in the remaining portion of grazed leaves, a net reduction in photosynthesis per leaf occurred only when the proportion of leaf area grazed exceeded 40% for Alnus incana and 23% for Alnus glutinosa. Since vein perforation by Agelastica alni was observed much more frequently in leaves of Betula than in Alnus, we hypothesized that declining Asat in herbivorized Betula was related to this disruption of water transport. A field experiment with artificial leaf perforation demonstrated a greater decline in Asat in vein-perforated Betula leaves than perforated leaves with midrib veins intact. However, regardless of leaf perforation regime, birch never showed post-perforation increases in Asat . In all species, rates of transpiration of grazed leaves linearly increased and water-use efficiency decreased with increased amount of leaf perforation. In grazed Alnus incana leaves, increasing leaf area consumption by Agelastica alni resulted in an increase of total phenols, a reduction in starch content and no changes in nitrogen concentration in the remaining portion. The increase in photosynthesis in Alnus incana might be related to declining leaf starch concentration or increasing stomatal conductance, but was unrelated to leaf nitrogen concentration. These gas exchange and leaf chemistry measurements suggest that in contrast to B. pendula, Alnus incana and Alnus glutinosa, which are the major host species for Agelastica alni, possess leaf-level physiological adaptations and defence mechanisms which can attenuate negative effects of herbivory by the alder leaf-beetle.

Relationship of aluminium and calcium to net CO2 exchange among diverse Scots pine provenances under pollution stress in Poland.

Light-saturated net photosynthesis (Asat), dark respiration (RD), and foliar nutrient content of eight European Scots pine (Pinus sylvestris L.) provenances were measured at experimental sites in western Poland. Two-year-old seedlings were planted in 1984 at two sites with similar soils in areas of contrasting air pollution. One site was near a point source of SO2 and other pollutants, and another 12 km to the southeast in an area free of acute air pollution was treated as a control. The eight provenances were from a large north-tosouth latitudinal range (60 to 43° N). At the heavily polluted site Scots pine trees exhibited lower growth rates and crown dieback and deformation. Soil pH, Ca and Mg were at least 10 times lower, and Al 10 times higher at the polluted than the control site. In 1991, concentrations of Al, P, Ca, S, Mn, Fe, and Zn in oneyear old Scots pine foliage were higher and Mg lower at the polluted than control site. At both sites foliar Mg levels were within the range considered deficient (≤0.6 mg g-1), and at the polluted site, Al concentrations were very high (670 to 880 µg g-1). In all provenances, RD of one-year-old needles was higher (by 22% on average) and Asat was lower (by 37% on average) at the polluted than the control site. The ratio of Asat: RD was half as great in all provenances at the polluted (4 to 6) than control site (8 to 11). Provenances of southern origin had greater increases in RD and water-use efficiency at the polluted site than other provenances. Within the polluted site alone, or across both sites, Asat in Scots pine was negatively correlated to the Al: Ca ratio (p<0.001, r=-0.93). Across sites RD increased with needle N and Al (multiple regression, p<0.001). The data suggest that at the polluted site there is excessive soil Al and deficient Mg availability, low needle Mg and high Al concentrations and high Al: Ca ratios, and that these have resulted in reduced photosynthetic capacity and increased respiration.

Height growth of different European Scots pine Pinus sylvestris L. Provenances in a heavily polluted and a control environment.

Results are presented of height measurements and degree of needle injury on five-year-old plants of Scots pine (Pinus sylvestris L.) growing near a phosphate fertiliser plant that emits SO(2) and fluorides. The populations of Scots pine represented in this experiment originate from 11 countries and were substantially differentiated in height growth and extent of needle necroses. Those populations which grew most rapidly were found to be the most sensitive to pollutant injury. The least productive provenances from the north of the range (Sweden, USSR) are at the same time characterized by lowest decline in height growth, lowest mortality and least extensive necroses. It is proposed that gene banks be established for the best genotypes likely to be eliminated in the heavily polluted conditions of Poland today.

Fungal diversity of Norway spruce litter: effects of site conditions and premature leaf fall caused by bark beetle outbreak.

Fungi play an important role in leaf litter decomposition due to their ability to break down the lignocellulose matrix, which other organisms are unable to digest. However, little is known regarding the factors affecting components of fungal diversity. Here, we quantified richness of internal fungi in relation to litter nutrient and phenolic concentrations, sampling season (spring or fall), and premature leaf shedding due to low precipitation and infestation of bark beetles (mainly Ips typographus and Ips duplicatus). The study was conducted in 37-year-old Norway spruce [Picea abies (L.) Karst.] stands, with three plots each in mixed forest (MF) and coniferous forest (CF) site conditions in south-central Poland. Fifty-four species of sporulating fungi were identified in 2,330 freshly fallen needles sampled during 2003-2005, including 45 species in MF and 31 in CF. The significantly higher number of species in MF was likely related to moister conditions at that site. Among isolated fungi, 22% (12 species) were identified as endophytes of Norway spruce in prior studies. During spring of 2005, we found less than half the number of isolates and fungal species at each forest site as compared to fall for the two prior years. This pattern was observed in typical soil fungi (e.g., Penicillium daleae, Penicillium purpurogenum) and endophytes/epiphytes (e.g., Aureobasidium pullulans, Alternaria alternata, Cladosporium spp., and Lophodermium piceae). Premature shedding of needles was the most likely cause of this decline because it shortened the time period for fungi to infect green needles while on the tree. For all sites and sampling periods, richness of internal fungi was strongly and positively related to the age of freshly fallen litter (assessed using needle Ca concentration as a needle age tracer) and was also negatively related to litter phenolic concentration. Richness of internal fungi in freshly fallen litter may be adversely affected by low soil moisture status, natural inhibitors slowing fungal colonization (e.g., phenolics) and biotic (e.g., insect infestation) and abiotic (e.g., drought) factors that shorten leaf life span.

Soil modification by different tree species influences the extent of seedling ectomycorrhizal infection.

Established vegetation can facilitate the ectomycorrhizal infection of seedlings, but it is not known whether this interaction is limited by the phylogenetic relatedness of trees and seedlings. We use a series of bioassay experiments to test whether soil modification by different ectomycorrhizal tree species causes different levels of seedling infection, whether the extent of seedling infection is a function of the relatedness of tree and seedling, and whether the effect of trees on seedlings is mediated by biotic or abiotic soil factors. We found that soils from under different tree species do vary in their mycorrhizal infectiveness. However, this variation is not related to the genetic relatedness of trees and seedlings but instead, appears to be an attribute of the overstory species, irrespective of seedling species, mediated through a suite of humus- and base-cation-related abiotic effects on soils. Modification of abiotic soil properties by overstory trees should be considered as an important factor in the effect of different overstory trees on the extent of seedling mycorrhizal infection.

Seedlings of five boreal tree species differ in acclimation of net photosynthesis to elevated CO(2) and temperature.

Biochemical models of photosynthesis suggest that rising temperatures will increase rates of net carbon dioxide assimilation and enhance plant responses to increasing atmospheric concentrations of CO(2). We tested this hypothesis by evaluating acclimation and ontogenetic drift in net photosynthesis in seedlings of five boreal tree species grown at 370 and 580 &mgr;mol mol(-1) CO(2) in combination with day/night temperatures of 18/12, 21/15, 24/18, 27/21, and 30/24 degrees C. Leaf-area-based rates of net photosynthesis increased between 13 and 36% among species in plants grown and measured in elevated CO(2) compared to ambient CO(2). These CO(2)-induced increases in net photosynthesis were greater for slower-growing Picea mariana (Mill.) B.S.P., Pinus banksiana Lamb., and Larix laricina (Du Roi) K. Koch than for faster-growing Populus tremuloides Michx. and Betula papyrifera Marsh., paralleling longer-term growth differences between CO(2) treatments. Measures at common CO(2) concentrations revealed that net photosynthesis was down-regulated in plants grown at elevated CO(2). In situ leaf gas exchange rates varied minimally across temperature treatments and, contrary to predictions, increasing growth temperatures did not enhance the response of net photosynthesis to elevated CO(2) in four of the five species. Overall, the species exhibited declines in specific leaf area and leaf nitrogen concentration, and increases in total nonstructural carbohydrates in response to CO(2) enrichment. Consequently, the elevated CO(2) treatment enhanced rates of net photosynthesis much more when expressed on a leaf area basis (25%) than when expressed on a leaf mass basis (10%). In all species, rates of leaf net CO(2) exchange exhibited modest declines with increasing plant size through ontogeny. Among the conifers, enhancements of photosynthetic rates in elevated CO(2) were sustained through time across a wide range of plant sizes. In contrast, for Populus tremuloides and B. papyrifera, mass-based photosynthetic rates did not differ between CO(2) treatments. Overall, net photosynthetic rates were highly correlated with relative growth rate as it varied among species and treatment combinations through time. We conclude that interspecific variation may be a more important determinant of photosynthetic response to CO(2) than temperature.

Nutrient conservation increases with latitude of origin in European Pinus sylvestris populations.

Nutrient availability varies across climatic gradients, yet intraspecific adaptation across such gradients in plant traits related to internal cycling and nutrient resorption remains poorly understood. We examined nutrient resorption among six Scots pine (Pinus sylvestris L.) populations of wide-ranging origin grown under common-garden conditions in Poland. These results were compared with mass-based needle N and P for 195 Scots pine stands throughout the species' European range. At the common site, green needle N (r(2)=0.81, P=0.01) and P (r(2)=0.58, P=0.08) concentration increased with increasing latitude of population origin. Resorption efficiency (the proportion of the leaf nutrient pool resorbed during senescence) of N and P of Scots pine populations increased with the latitude of seed origin (r(2) > or = 0.67, P < or = 0.05). The greater resorption efficiency of more northerly populations led to lower concentrations of N and P in senescent leaves (higher resorption proficiency) than populations originating from low latitudes. The direction of change in these traits indicates potential adaptation of populations from northern, colder habitats to more efficient internal nutrient cycling. For native Scots pine stands, results showed greater nutrient conservation in situ in cold-adapted northern populations, via extended needle longevity (from 2 to 3 years at 50 degrees N to 7 years at 70 degrees N), and greater resorption efficiency and proficiency, with their greater resorption efficiency and proficiency having genotypic roots demonstrated in the common-garden experiment. However, for native Scots pine stands, green needle N decreased with increasing latitude (r(2)=0.83, P=0.0002), and P was stable other than decreasing above 62 degrees N. Hence, the genotypic tendency towards maintenance of higher nutrient concentrations in green foliage and effective nutrient resorption, demonstrated by northern populations in the common garden, did not entirely compensate for presumed nutrient availability limitations along the in situ latitudinal temperature gradient.

Evidence that longer needle retention of spruce and pine populations at high elevations and high latitudes is largely a phenotypic response.

There is abundant evidence that evergreen conifers living at high elevations or at high latitudes have longer-lived needles than trees of the same species living elsewhere. This pattern is likely caused by the influence of low temperature in combination with related factors such as a short growing season and low nutrient availability. Because it is not known to what degree such patterns result from phenotypic versus genotypic variation, we evaluated needle longevity for common-garden-grown lowland populations of European Scots pine (Pinus sylvestris L.) of wide latitudinal origin and Norway spruce (Picea abies L.) of wide elevational origin. Nine-year-old trees of 16 Scots pine populations ranging in origin from 47 degrees to 60 degrees N were studied in Kórnik, Poland (52 degrees N) and 18-year-old trees of 18 Norway spruce populations ranging in origin from 670 to 1235 m elevation in southwestern Poland were studied near Morawina, Poland (51 degrees N, 180 m elevation). There was no tendency in either species for populations from northern or high elevation origins to retain needles longer than other populations. All of the Scots pine populations had between 2.5 to 3.0 needle age cohorts and all of the Norway spruce populations had between 6.4 and 7.2 needle age cohorts. Thus, extended needle retention in Scots pine and Norway spruce populations in low-temperature habitats at high elevations and high latitudes appears to be largely an environmentally regulated phenotypic acclimation.

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.