Piotr Karolewski

TREE SPECIES EFFECTS ON DECOMPOSITION AND FOREST FLOOR DYNAMICS IN A COMMON GARDEN

We studied the effects of tree species on leaf litter decomposition and forest floor dynamics in a common garden experiment of 14 tree species (Abies alba, Acer platanoides, Acer pseudoplatanus, Betula pendula, Carpinus betulus, Fagus sylvatica, Larix decidua, Picea abies, Pinus nigra, Pinus sylvestris, Pseudotsuga menziesii, Quercus robur, Quercus rubra, and Tilia cordata) in southwestern Poland. We used three simultaneous litter bag experiments to tease apart species effects on decomposition via leaf litter chemistry vs. effects on the decomposition environment. Decomposition rates of litter in its plot of origin were negatively correlated with litter lignin and positively correlated with mean annual soil temperature (MAT(soil)) across species. Likewise, decomposition of a common litter type across all plots was positively associated with MAT(soil), and decomposition of litter from all plots in a common plot was negatively related to litter lignin but positively related to litter Ca. Taken together, these results indicate that tree species influenced microbial decomposition primarily via differences in litter lignin (and secondarily, via differences in litter Ca), with high-lignin (and low-Ca) species decomposing most slowly, and by affecting MAT(soil), with warmer plots exhibiting more rapid decomposition. In addition to litter bag experiments, we examined forest floor dynamics in each plot by mass balance, since earthworms were a known component of these forest stands and their access to litter in litter bags was limited. Forest floor removal rates estimated from mass balance were positively related to leaf litter Ca (and unrelated to decay rates obtained using litter bags). Litter Ca, in turn, was positively related to the abundance of earthworms, particularly Lumbricus terrestris. Thus, while species influence microbially mediated decomposition primarily through differences in litter lignin, differences among species in litter Ca are most important in determining species effects on forest floor leaf litter dynamics among these 14 tree species, apparently because of the influence of litter Ca on earthworm activity. The overall influence of these tree species on leaf litter decomposition via effects on both microbial and faunal processing will only become clear when we can quantify the decay dynamics of litter that is translocated belowground by earthworms.

Foliage Age and Pollution Alter Content of Phenolic Compounds and Chemical Elements in Pinus nigra Needles

Changes of phenolics and chemical elements [nitrogen (N), phosphorus (P), potassium (K), calcium (Ca), magnesium (Mg), manganese (Mn), iron (Fe), boron (B), copper (Cu), zinc (Zn), lead (Pb), cadmium (Cd), chromium (Cr), nickel (Ni) and cobalt (Co)] content in needles of black pine ( Pinus nigra) as dependent on age of needles (5–6 classes) and pollution were examined. The content of ortho-diphenols (o-dPh) and total phenols (TPh) was significantly higher at a polluted site than at a control one. It increased with age of needles at both sites. At the polluted site contents of N, K, Mg in black pine needles were lower and of Fe, Ni and F were higher than at the control site. An increase of content with age of needles at both sites was detected for Ca, Fe, B and F, and a decrease for N, P, K, Cu and Ni. The content of elements in different age classes of needles is connected with their mobility. The content of phenolics is negatively correlated with main nutrients and positively with some toxic elements.

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 (Nmass) 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 (Narea). In evergreen conifers, LMA and Narea were less powerful predictors of area-based photosynthetic rate (Amax(area)) in comparison with deciduous broadleaved angiosperms. Multiple regression for sun and shade leaves showed that, in each group, Amax(mass) was related to Nmass but not to LMA, whereas LMA became a significant codeterminant of Amax(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.

Ontogenetic patterns of leaf CO2 exchange, morphology and chemistry in Betula pendula trees

Abstract In order to explore ontogenetic variation in leaf-level physiological traits of Betula pendula trees, we measured changes in mass- (Amass) and area-based (Aarea) net photosynthesis under light-saturated conditions, mass- (RSmass) and area-based (RSarea) leaf respiration, relative growth rate, leaf mass per area (LMA), total nonstructural carbohydrates (TNC), and macro- and micronutrient concentrations. Expanding leaves maintained high rates of Aarea, but due to high growth respiration rates, net CO2 fixation occurred only at irradiances >200 µmol photons m–2 s–1. We found that full structural leaf development is not a necessary prerequisite for maintaining positive CO2 balance in young birch leaves. Maximum rates of Aarea were realized in late June and early July, whereas the highest values of Amass occurred in May and steadily declined thereafter. The maintenance respiration rate averaged ≈8 nmol CO2 g–1 s–1, whereas growth respiration varied between 0 and 65 nmol CO2 g–1 s–1. After reaching its lowest point in mid-June, leaf respiration increased gradually until the end of the growing season. Mass and area-based dark respiration were significantly positively correlated with LMA at stages of leaf maturity, and senescence. Concentrations of P and K decreased during leaf development and stabilized or increased during maturity, and concentrations of immobile elements such as Ca, Mn and B increased throughout the growing season. Identification of interrelations between leaf development, CO2 exchange, TNC and leaf nutrients allowed us to define factors related to ontogenetic variation in leaf-level physiological traits and can be helpful in establishing periods appropriate for sampling birch leaves for diagnostic purposes such as assessment of plant and site productivity or effects of biotic or abiotic factors.

Altered root growth and plant chemistry of Pinus sylvestris seedlings subjected to aluminum in nutrient solution

One-year-old Scots pine (Pinus sylvestris L.) seedlings were grown for 9 weeks in nutrient solutions containing 0, 0.5, 1, 2 and 4 mM aluminum nitrate (Al(NO3)3) at pH 4.2. Nine weeks exposure to Al significantly reduced total plant, shoot and root mass and caused a linear decline in proportional allocation of biomass to roots. Relative growth rate of roots declined to as low as zero. Aluminum treatment decreased calcium and magnesium uptake and increased Al content in roots and needles. After 3 weeks of exposure a 10–60% increase in total phenols in roots and a 20–40% increase in o-diphenols in roots and needles were noted. Roots affected by Al showed degeneration of meristematic cells, fewer cell divisions, deformation in cell walls and higher lignification and suberization. The majority of root apices were structurally similar to dormant roots, and a premature senescence of the entire root system was observed. Net photosynthetic rate after 6 weeks of treatment was negatively correlated with needle Al content and Al/Ca ratio (r < -0.9, P < 0.1). The results suggest that Scots pine may be more susceptible to Al than was expected based on previous experiments.

Needle CO2 exchange, structure and defense traits in relation to needle age in Pinus heldreichii Christ - A relict of Tertiary flora

Abstract Pinus heldreichii Christ is a long-lived, slow-growing Tertiary relict from the Balkans. In this study we evaluated the physiological characteristics of eight needle-age classes of P. heldreichii grown at the Arboretum of the Institute of Dendrology in Kórnik, Poland. At the end of the growing season, current-year foliage had the highest rates of mass-based light-saturated net photosynthesis (Asat) of 33.5 nmol CO2· g–1· s–1. Asat decreased with needle age, but older needle classes retained from approximately 62 to 26% of the current needles’ rate. The relationship between leaf N and chlorophyll a concentration among all needle-age classes was highly significant (r = 0.96, P = 0.0006). The variation in Asat of 1- to 7-year-old needles was linearly related to needle N concentration (r = 0.98, P = 0.0001). Needle dark respiration rates among these needle age classes ranged from 0.8 to 2.2 nmol · g–1· s–1 and decreased with needle age and nitrogen concentration. Total phenols and glucose concentrations increased linearly with needle age. A similar pattern was observed in acid buffering capacity and the pH of tissue homogenates. The water content ranged from 62% for the current needles to 51% for the 6-year-old needles. Greater investment in leaf structural tissue and increased chemical defense is associated with higher structural cost of older needles and may reduce their photosynthetic activity. Significant declines in water and nitrogen content with needle age and an increase in content of phenolics is most likely a defense adaptation of P. heldreichii related to the species’ long-lived leaves.

Phenotypic correlates of the lianescent growth form: a review

Background As proposed by Darwin, climbers have been assumed to allocate a smaller fraction of biomass to support organs in comparison with self-supporting plants. They have also been hypothesized to possess a set of traits associated with fast growth, resource uptake and high productivity. Scope In this review, these hypotheses are evaluated by assembling and synthesizing published and unpublished data sets from across the globe concerning resource allocation, growth rates and traits of leaves, stems and roots of climbers and self-supporting species. Conclusions The majority of studies offer little support for the smaller allocation of biomass to stems or greater relative growth rates in climbers; however, these results are based on small sized (<1 kg) plants. Simulations based on allometric biomass equations demonstrate, however, that larger lianas allocate a greater fraction of above-ground biomass to leaves (and therefore less biomass to stems) compared with similar sized trees. A survey of leaf traits of lianas revealed their lower average leaf mass per area (LMA), higher N and P concentration and a slightly higher mass-based photosynthetic rate, as well as a lower concentration of phenolic-based compounds than in woody self-supporting species, consistent with the specialization of lianas towards the fast metabolism/rapid turnover end of the global trait spectra. Liana stems have an efficient hydraulic design and unique mechanical features, while roots appear to penetrate deeper soil levels than in trees and are often able to generate hydraulic pressure. Much remains to be learned, however, about these and other functional specializations of their axial organs and the associated trade-offs. Developmental switches between self-supporting, searcher and climbing shoots within the same individual are a promising field of comparative studies on trait association in lianas. Finally, some of the vast trait variability within lianas may be reduced when species with different climbing mechanisms are considered separately, and when phylogenetic conservatism is accounted for.

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.

Influence of mineral fertilization on food quality of oak leaves and utilization efficiency of food components by the gypsy moth

Host plant quality is a key determinant of the performance of larvae of herbivorous insects. The effects of nitrogen and dolomite fertilization on the quality of pedunculate oak, Quercus robur L. (Fagaceae) foliage, as a food for gypsy moth, Lymantria dispar L. (Lepidoptera: Lymantriidae) larvae were evaluated. The seedlings were divided into five fertilization treatments (nonfertilized control, commercial nutrient solution, commercial nutrient solution + (NH4)2SO4, commercial nutrient solution + KNO3, and commercial nutrient solution + dolomite). The experiment was performed in Petri dishes, in each of which a fresh leaf from one treatment and one larva were placed. Insect performance assays, survival, development, growth, and food utilization were evaluated for each fertilization treatment. Leaf samples were assayed for nitrogen and other main nutrients, soluble carbohydrates, and phenolic compounds. The fertilizer treatment with added ammonium improved gypsy moth performance, and the amount of food eaten was the lowest in this treatment. Utilization of elements from the food depended on the element and on the fertilization treatment. The insect bodies retained 50–64% of the nitrogen and 55–79% of the phosphorus. The results show that the efficiency of conversion of ingested food (ECI) and the efficiency of conversion of digested food (ECD) differ among the fertilization treatments, but it is not possible to define a general trend. Our results suggest that fertilization (especially ammonium) of host plants can increase herbivore performance, decrease the amount of food needed, and increase its utilization efficiency.

Variation in aboveground net primary production of diverse European Pinus sylvestris populations

Abstract On range-wide and regional scales, climate and site factors exert control over tree growth, masking the genetic basis of biomass accumulation and allocation. To determine intrinsic population differences in productivity, aboveground net primary production (ANPP) was measured in 16-year-old Scots pine from 19 geographically distinct populations grown in a common garden experiment in central Poland (52°N). The populations originated from the northern (>55°N), central (54–47°N), and southern (<45°N) European range of Scots pine. We calculated ANPP from aboveground growth components, using diameter-based allometric equations developed for this site. Average foliage, aboveground woody and total ANPP differed significantly among populations and were greater for central European populations than for the southern and northern ones. Stocking and total ANPP per tree were positively correlated to stand aboveground biomass (r2≥0.71). The relationship between the latitude of seed origin and ANPP was curvilinear and maximum for populations originating near the planting site (52°N). ANPP declined in populations with increasing longitude eastward from the Atlantic Ocean towards the center of the continent. This study underscores the potentially large genetic control of ANPP and biomass accumulation among diverse Scots pine populations.