Are correlations among foliar traits in ferns consistent with those in the seed plants?

Broad-based studies of gymnosperms and angiosperms reveal consistent and functionally significant correlations among foliar traits such as leaf mass per area (LMA), maximum photosynthetic rate (A(area)), foliar nitrogen (N(area)), foliar chlorophyll (Chl) and leaf longevity. To assess the generality of these relationships, we studied 20 fern species growing in the understorey of a temperate deciduous forest. We found that foliar N(area) increases with LMA, and that foliar N(area) and A(area) are positively correlated with one another, as are foliar N(area) and Chl. The ferns in general have very low LMA compared with most seed plants; A(area), N(area) and Chl are below median values for seed plants but are not extreme. Species with overwintering fronds have significantly higher LMA than species with fronds that senesce at the end of the growing season, as well as a significantly higher C : N ratio in frond tissue and relatively high foliar N on an areal basis. Correlations among foliar traits associated with gas exchange in these forest understorey ferns are in accordance with patterns reported for seed plants, suggesting a high degree of functional constraint on the interrelationships among key elements in foliar design.     

Nitrogen resorption and photosynthetic activity over leaf life span in an evergreen shrub, Rhododendron ferrugineum, in a subalpine environment

Here, the advantages for a shrub of having long vs short-lived leaves was investigated in Rhododendron ferrugineum by following nitrogen(15N) and carbon(14C) resorption and translocation, and photosynthetic capacity over the life span. Mean leaf life span was 19 months. Nitrogen (N) resorption in attached leaves occurred mainly in the first year (23%) and reached a maximum of 31% in the second. Although, resorption was similar in attached and fallen 1-yr-old leaves, it was on average one-third higher in fallen than in attached older leaves. Final N resorption of a leaf compartment reached 41%, half occurring from healthy leaves during the first year. Photosynthetic capacity decreased slightly during the life span. Before shoot growth, plant photosynthesis was mainly supported by 1-yr-old leaves, although the contribution of the 2-yr-old leaves was nonnegligible (15% of the capacity and higher carbon transfer toward the roots). After shoot growth, the current-year leaves made the greatest contribution. Our results suggest that short-lived leaves (half of the cohort) are mainly involved in a photosynthetic function, having a high photosynthetic capacity and drawing most of their resorbed N towards current-year leaves; and long-lived leaves are also involved in a conservative function, increasing N resorption and mean residence time (MRT).     

Photosynthetic stimulation under long-term CO2 enrichment and fertilization is sustained across a closed Populus canopy profile (EUROFACE)

The long-term response of leaf photosynthesis to rising CO2 concentrations [CO2] depends on biochemical and morphological feedbacks. Additionally, responses to elevated [CO2] might depend on the nutrient availability and the light environment, affecting the net carbon uptake of a forest stand. After 6 yr of exposure to free-air CO2 enrichment (EUROFACE) during two rotation cycles (with fertilization during the second cycle), profiles of light, leaf characteristics and photosynthetic parameters were measured in the closed canopy of a poplar (Populus) short-rotation coppice. Net photosynthetic rate (A(growth)) was 49% higher in poplars grown in elevated [CO2], independently of the canopy position. Jmax significantly increased (15%), whereas leaf carboxylation capacity (Vcmax), leaf nitrogen (N(a)) and chlorophyll (Chl(a)) were unaffected in elevated [CO2]. Leaf mass per unit area (LMA) increased in the upper canopy. Fertilization created more leaves in the top of the crown. These results suggest that the photosynthetic stimulation by elevated [CO2] in a closed-canopy poplar coppice might be sustained in the long term. The absence of any down-regulation, given a sufficient sink capacity and nutrient availability, provides more carbon for growth and storage in this bioenergy plantation.     

Leaf phenology and seasonal variation of photosynthesis of invasive <Emphasis Type="Italic">Berberis thunbergii</Emphasis> (Japanese barberry) and two co-occurring native understory shrubs in a northeastern United States deciduous forest

Early leafing and extended leaf longevity can be important mechanisms for the invasion of the forest understory. We compared the leaf phenology and photosynthetic characteristics of Berberis thunbergii, an early leafing invasive shrub, and two co-occurring native species, evergreen Kalmia latifolia and late leafing Vaccinium corymbosum, throughout the 2004 growing season. Berberis thunbergii leafed out 1 month earlier than V. corymbosum and approximately 2 weeks prior to the overstory trees. The photosynthetic capacity [characterized by the maximum carboxylation rate of Rubisco (V _cmax) and the RuBP regeneration capacity mediated by the maximum electron transport rate (J _max)] of B. thunbergii was highest in the spring open canopy, and declined with canopy closure. The 2003 overwintering leaves of K. latifolia displayed high V _cmax and J _max in spring 2004. In new leaves of K. latifolia produced in 2004, the photosynthetic capacity gradually increased to a peak in mid-September, and reduced in late November. V. corymbosum, by contrast, maintained low V _cmax and J _max throughout the growing season. In B. thunbergii, light acclimation was mediated by adjustment in both leaf mass per unit area and leaf N on a mass basis, but this adjustment was weaker or absent in K. latifolia and V. corymbosum. These results indicated that B. thunbergii utilized high irradiance in the spring while K. latifolia took advantage of high irradiance in the fall and the following spring. By contrast, V. corymbosum generally did not experience a high irradiance environment and was adapted to the low irradiance understory. The apparent success of B. thunbergii therefore, appeared related to a high spring C subsidy and subsequent acclimation to varying irradiance through active N reallocation and leaf morphological modifications.     

Is microbial community composition in boreal forest soils determined by pH,C-to-N ratio,the trees,or all three?

In Fennoscandian boreal forests, soil pH and N supply generally increase downhill as a result of water transport of base cations and N, respectively. Simultaneously, forest productivity increases, the understory changes from ericaceous dwarf shrubs to tall herbs; in the soil, fungi decrease whereas bacteria increase. The composition of the soil microbial community is mainly thought to be controlled by the pH and C-to-N ratio of the substrate. However, the latter also determines the N supply to plants, the plant community composition, and should also affect plant allocation of C below ground to roots and a major functional group of microbes, mycorrhizal fungi. We used phospholipid fatty acids (PLFAs) to analyze the potential importance of mycorrhizal fungi by comparing the microbial community composition in a tree-girdling experiment, where tree belowground C allocation was terminated, and in a long-term (34 years) N loading experiment, with the shifts across a natural pH and N supply gradient. Both tree girdling and N loading caused a decline of ca. 45% of the fungal biomarker PLFA 18:2ω6,9, suggesting a common mechanism, i.e., that N loading caused a decrease in the C supply to ectomycorrhizal fungi just as tree girdling did. The total abundance of bacterial PLFAs did not respond to tree girdling or to N loading, in which cases the pH (of the mor layer) did not change appreciably, but bacterial PLFAs increased considerably when pH increased across the natural gradient. Fungal biomass was high only in acid soil (pH < 4.1) with a high C-to-N ratio (>38). According to a principal component analysis, the soil C-to-N ratio was as good as predictor of microbial community structure as pH. Our study thus indicated the soil C-to-N ratio, and the response of trees to this ratio, as important factors that together with soil pH influence soil microbial community composition.     

Wide geographical and ecological distribution of nitrogen and carbon gains from fungi in pyroloids and monotropoids (Ericaceae) and in orchids

* Stable isotope abundance analyses recently revealed that some European green orchids and pyroloids (Ericaceae) are partially myco-heterotrophic, exploiting mycorrhizal fungi for organic carbon and nitrogen. Here we investigate related species to assess their nutritional mode across various forest and climate types in Germany and California. * C- and N-isotope signatures of five green pyroloids, three green orchids and several obligate myco-heterotrophic species (including the putatively fully myco-heterotrophic Pyrola aphylla) were analysed to quantify the green plants' nutrient gain from their fungal partners and to investigate the constancy of enrichment in (13)C and (15)N of fully myco-heterotrophic plants from diverse taxa and locations relative to neighbouring autotrophic plants. * All green pyroloid and one orchid species showed significant (15)N enrichment, confirming incorporation of fungi-derived N compounds while heterotrophic C gain was detected only under low irradiance in Orthilia secunda. Pyrola aphylla had an isotope signature equivalent to those of fully myco-heterotrophic plants. * It is demonstrated that primarily N gain from mycorrhizal fungi occurred in all taxonomic groups investigated across a wide range of geographical and ecological contexts. The (13)C and (15)N enrichment of obligate myco-heterotrophic plants relative to accompanying autotrophic plants turned out as a fairly constant parameter.     

Nitrogen deposition but not ozone affects productivity and community composition of subalpine grassland after 3 yr of treatment

A field experiment was established at 2000 m above sea level (asl) in the central Swiss Alps with the aim of investigating the effects of elevated ozone (O(3)) and nitrogen deposition (N), and of their combination, on above-ground productivity and species composition of subalpine grassland. One hundred and eighty monoliths were extracted from a species-rich Geo-Montani-Nardetum pasture and exposed in a free-air O(3)-fumigation system to one of three concentrations of O(3) (ambient, 1.2 x ambient, 1.6 x ambient) and five concentrations of additional N. Above-ground biomass, proportion of functional groups and normalized difference vegetation index (NDVI) were measured annually. After 3 yr of treatment, the vegetation responded to the N input with an increase in above-ground productivity and altered species composition, but without changes resulting from elevated O(3). N input > 10 kg N ha(-1) yr(-1) was sufficient to affect the composition of functional groups, with sedges benefiting over-proportionally. No interaction of O(3) x N was observed, except for NDVI; positive effects of N addition on canopy greenness were counteracted by accelerated leaf senescence in the highest O(3) treatment. The results suggest that effects of elevated O(3) on the productivity and floristic composition of subalpine grassland may develop slowly, regardless of the sensitive response to increasing N.     

Litterfall and fine root biomass contribution to nutrient dynamics in second- and old-growth Douglas-fir ecosystems

Litterfall and fine root production were measured for three years as part of a carbon balance study of three forest stands in the Pacific Northwest of the United States. A young second-growth Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] stand, a second-growth Douglas-fir with red alder (Alnus rubra Bong.) stand, and an old-growth (∼550 years) Douglas-fir stand were monitored for inputs of carbon and nitrogen into the soil from litterfall and fine root production, as well as changes in soil C and N. Fine root production and soil nutrient changes were measured through the use of soil ingrowth bags containing homogenized soil from the respective stands. Litterfall biomass was greatest in the Douglas-fir-alder stand (527 g m⁻² yr⁻¹) that annually returned nearly three times the amount of N as the other stands. Mean residence time for forest floor material was also shortest at this site averaging 4.6 years and 5.5 years for C an N, respectively. Fine root production in the upper 20 cm ranged from 584 g m⁻² in the N rich Douglas-fir-alder stand to 836 g m⁻² in the old-growth stand. Fine root production (down to one meter) was always greater than litterfall with a below:above ratio ranging from 3.73 for the young Douglas-fir stand to 1.62 for the Douglas-fir-alder stand. The below:above N ratios for all three stands closely approximate those for biomass. Soil changes in both C and N differed by site, but the soil C changes in the old-growth stand mirrored those obtained in an ongoing CO₂ flux study. Results from the soil ingrowth bags strongly suggest that this method provides a simple, but sufficient device for measuring potential fine root biomass production as well as soil chemical changes.     

Competition for nitrogen between Pinus sylvestris and ectomycorrhizal fungi generates potential for negative feedback under elevated CO2

We investigated fungal species-specific responses of ectomycorrhizal (ECM) Scots pine (Pinus sylvestris) seedlings on growth and nutrient acquisition together with mycelial development under ambient and elevated CO₂. Each seedling was associated with one of the following ECM species: Hebeloma cylindrosporum, Laccaria bicolor, Suillus bovinus, S. luteus, Piloderma croceum, Paxillus involutus, Boletus badius, or non-mycorrhizal, under ambient, and elevated CO₂ (350 or 700 μl l⁻¹ CO₂); each treatment contained six replicates. The trial lasted 156 days. During the final 28 days, the seedlings were labeled with ¹⁴CO₂. We measured hyphal length, plant biomass, ¹⁴C allocation, and plant nitrogen and phosphorus concentration. Almost all parameters were significantly affected by fungal species and/or CO₂. There were very few significant interactions. Elevated CO₂ decreased shoot-to-root ratio, most strongly so in species with the largest extraradical mycelium. Under elevated CO₂, ECM root growth increased significantly more than hyphal growth. Extraradical hyphal length was significantly negatively correlated with shoot biomass, shoot N content, and total plant N uptake. Root dry weight was significantly negatively correlated with root N and P concentration. Fungal sink strength for N strongly affected plant growth through N immobilization. Mycorrhizal fungal-induced progressive nitrogen limitation (PNL) has the potential to generate negative feedback with plant growth under elevated CO₂. Responsible Editor: Herbert Johannes Kronzucker