Effects of soil water on photosynthetic characteristics and leaf traits of Cyclobalanopsis glauca seedlings growing under nutrient-rich and -poor soil

Cyclobalanopsis glauca is a dominant species in mid-subtropical forest, and usually plays an important role in forest ecosystems. However, it often suffers redundant precipitation or water stress, which often concurs with high temperature, nutrient depletion and strong irradiance. The study presented in the paper hypothesized that soil water exerted strong influence on leaf gas exchange and traits. The objective of this study was to clarify the effect of soil water changes on photosynthetic characteristics and leaf traits and their relationships of C. glauca seedlings growing on nutrient-rich and nutrient-poor soil at three water levels. The study measured the specific leaf area (SLA), nitrogen content, chlorophyll concentrations and photosynthetic light response curve. Its results showed that there were no differences in leaf size, leaf dry weight, SLA, leaf dry matter content, Leaf nitrogen concentration and Leaf chlorophyll between the two soil nutrient treatments, while these parameters differed significantly among different water levels for either of the treatments. There were large variations in leaf photosynthetic parameters and leaf traits among the different water treatments, indicating different response patterns of C. glauca seedling and its adaptation to the different soil water conditions. There were no significant differences in light-saturated photosynthetic rate (A_max) and apparent quantum yield (Ø) between the nutrient-rich and nutrient-poor soils, which indicated that the C. glauca seedlings could maintain similar capacities in different soils that differed in nutrient condition. As to the relation between the photosynthesis and leaf traits, the A_max and PNUE were positively correlated with the SLA, respectively, but the SLA had significant negative relationship with the leaf N (P<0.01) in nutrient-rich soil. In contrast, both A_max and PNUE were significantly negatively correlated with the SLA, respectively (P<0.01); and the SLA was not significantly positively correlated with the leaf nitrogen concentration of the nutrient-poor soil (P>0.05). The specific leaf areas (SLA), nitrogen and chlorophyll concentrations as well as other photosynthetic features were influenced in a coordinative manner by the soil water. The relation among the A_max, PNUE and the N_mass, SLA could be described as a binomial equation and a liner negative regression for the nutrient-rich and nutrient-poor soil, respectively. In conclusion, soil water was more constraining factor than the soil nutrient to the photosynthesis of C. glauca seedlings, nutrient-rich soil could offset some negative influence resulting from soil water deficit on LSP and LCP. Factors affecting the variations of photosynthetic characteristics and leaf traits of C. glauca seedlings differed between the nutrient-rich and nutrient-poor soils.     

Nitrate production in the rhizosphere of Plantago species

Summary The production of nitrate in an old established dune grassland soil and its uptake by plants was studied by comparing amounts of mineral nitrogen and numbers of nitrifying bacteria in the rhizosphere on the one hand, and on the other accumulated nitrate and levels of nitrate reductase (NaR) of individual plants of three Plantago species,i. e., P. major, P. lanceolata andP. coronopus. For these three Plantago species andP. media basal levels of NaR in the absence of nitrate were determined in plants grown in culture solutions. The basal NaR levels ofP. major andP. media (species occurring on nutrient-rich soils) were significantly higher than those ofP. lanceolata andP. coronopus (species found on nutrient-poor soils). NaR activity increased in the presence of nitrate and was suppressed by ammonium.From the numbers of nitrifying bacteria in the rhizosphere and NaR activity in the leaves it was concluded that nitrate was produced in the root environments of the three Plantago species and that the compound was taken up by the plants. NaR activities and numbers of nitrifying bacteria were higher for individuals ofP. major than for those ofP. lanceolata andP. coronopus. No correlation was found between the ammonium levels and the numbers of nitrifying bacteria in the soil, and no indications of inhibition of nitrifying bacteria in the rhizosphere were obtained. For individuals ofP. lanceolata a correlation was found between the numbers of nitrifying bacteria in the soil and NaR activity in the leaves. The results are discussed in relation to the ecological habitats of the three species.Grassland Species Research Group Publication No.38.     

Differences between European native and American invasive populations of Lythrum salicaria

Abstract. We compared the demographic characteristics of native central-European and invasive USA-populations of Lythrum salicaria growing in similar habitats. Based on the ‘Evolution of increased competitive ability’ (EICA) hypothesis, we predicted that shoot density, height, and biomass, fertility, and fecundity would be less in the European populations due to greater loss of plant material caused by exposure to control agents. Shoot density was significantly greater in most USA-populations, but shoot height and biomass were similar in invasive and native populations growing in similar habitats, especially in growing seasons with greater water availability. The number of fertile shoots was greater for invasive populations, except for those growing in sandy, nutrient-poor substrates, while percent fertility did not differ between populations growing under similar field conditions. Fecundity was also similar for populations growing in nutrient-poor and intermediate habitats, but was significantly greater in USA-populations (89–103 seeds per fruit), compared to European populations (58–64 seeds per fruit), growing in nutrient-rich habitats; seed predators were found in these European populations only. Log-linear analysis of transition frequency matrices showed that the growth of USA-populations is different from European populations, but that habitat effect was strong. Population dynamics were similar for populations growing in nutrient-poor habitats; location was of marginal importance only. USA-populations responded differently from European populations growing in intermediate and nutrient-rich habitats; differences were most pronounced between invasive (center of its North American distribution) and native populations growing in nutrient-rich habitats. The use of insect herbivores as biological control agents in North America will work best against L. salicaria populations growing in nutrient-rich habitats in the center of its invasive distribution, but will be less effective against populations growing in other habitats or portions of its range. Other factors, including nutrient- or water availability, and climate, may be as important as herbivory in affecting invasive populations; these factors interact to control L. salicaria in a more complex manner than thought previously.     

Refuse dumps from leaf-cutting ant nests reduce the intensity of above-ground competition among neighboring plants in a Patagonian steppe

In arid environments, the high availability of sunlight due to the scarcity of trees suggests that plant competition take place mainly belowground for water and nutrients. However, the occurrence of soil disturbances that increase nutrient availability and thereby promote plant growth may enhance shoot competition between neighboring plants. We conducted a greenhouse experiment to evaluate the influence of the enriched soil patches generated by the leaf-cutting ant, Acromyrmex lobicornis, on the performance of the alien forb Carduus thoermeri (Asteraceae) under different intraspecific competition scenarios. Our results showed that substrate type and competition scenario affected mainly aboveground plant growth. As expected, plants growing without neighbors and in nutrient-rich ant refuse dumps showed more aboveground biomass than plants growing with neighbors and in nutrient-poor steppe soils. However, aboveground competition was more intense in nutrient-poor substrates: plants under shoot and full competition growing in the nutrient-rich ant refuse dumps showed higher biomass than those growing on steppe soils. Belowground biomass was similar among focal plants growing under different substrate type. Our results support the traditional view that increments in resource availability reduce competition intensity. Moreover, the fact that seedlings in this sunny habitat mainly compete aboveground illustrates how limiting factors may be scale-dependent and change in importance as plants grow.     

Comparisons of nutrient recovery and specific leaf area variation betweenCarex lasiocarpa var.occultans andCarex thunbergii var.appendiculata with reference to nutrient conditions and shading byPhragmites australis

The distribution of two sedge species was studied in two mires which differ in abiotic environments and in distribution ofPhragmites australis. Carex lasiocarpa var.occultans dominated in nutrient-poor valley mire, andCarex thunbergii var.appendiculata dominated in nutrient-rich flood plain subject to water fluctuations.Phragmites australis grew well in nutrient-rich conditions. The distribution ofC. lasiocarpa showed a strong negative correlation withP. australis coverage, whereasC. thunbergii coverage was not affected byP. australis. The leaf area per dry leaf mass (specific leaf area: SLA) ofC. thunbergii increased with shading byP. australis, but that ofC. lasiocarpa was stable. The SLA flexibility ofC. thunbergii to light interception might enable this species to invadeP. australis patches in nutrient-rich environments. The residual nutrient ratio of nitrogen and phosphorus (the ratio of the residual nutrient content at the end of the growing season to peak nutrient content) in the vegetative ramet ofC. thunbergii was 1.7 times higher than that ofC. lasiocarpa. This low residual ratio may indicate effective nutrient recovery to storage organs. The effective nutrient recovery inC. lasiocarpa might enable this species to grow even in nutrient-poor environments. However, it may be difficult forC. lasiocarpa to expand its habitat to nutrient-rich areas whereP. australis dominates as it is not shade tolerant.     

Variation of soil and biomass carbon pools in beech forests across a precipitation gradient

Temperate forests have recently been identified as being continuing sinks for carbon even in their mature and senescent stages. However, modeling exercises indicate that a warmer and drier climate as predicted for parts of Central Europe may substantially alter the source/sink function of these economically important ecosystems. In a transect study with 14 mature European beech (Fagus sylvatica L.) forests growing on uniform geological substrate, we analyzed the influence of a large reduction of annual precipitation (970–520 mm yr^?1) on the carbon stocks in fast and slow pools, independent of the well‐known aging effect. We investigated the C storage in the organic L, F, H layers, the mineral soil to 100 cm, and in the biomass (stem, leaves, fine roots), and analyzed the dependence of these pools on precipitation. Soil organic carbon decreased by about 25% from stands with > 900 mm yr^?1 to those with < 600 mm yr^?1; while the carbon storage in beech stems slightly increased. Reduced precipitation affected the biomass C pool in particular in the fine root fraction but much less in the leaf biomass and stem fractions. Fine root turnover increased with a precipitation reduction, even though stand fine root biomass and SOC in the organic L, F, and H layers decreased. According to regression analyses, the C storage in the organic layers was mainly controlled by the size of the fine root C pool suggesting an important role of fine root turnover for the C transfer from tree biomass to the SOC pool. We conclude that the long‐term consequence of a substantial precipitation decrease would be a reduction of the mineral soil and organic layer SOC pools, mainly due to higher decomposition rates. This could turn temperate beech forests into significant carbon sources instead of sinks under global warming.     

Differential growth patterns and fitness may explain contrasted performances of the invasive <Emphasis Type="Italic">Prunus serotina</Emphasis> in its exotic range

This research investigates why the invasive American black cherry tends to dominate the forest canopy on well-drained, nutrient-poor soils, but usually hardly establishes on both waterlogged and calcareous soils in its exotic range. Prunus serotina was sampled from four soil types and two light conditions, to measure (1) radial growth; (2) height growth compared to the main native competitor, Fagus sylvatica; (3) leaf traits; (4) seed production; and (5) rate of fungal attack. We found that P. serotina invested a significant amount of energy in height growth and seed production on well-drained, nutrient-poor soils. These characteristics enabled it to rapidly capture canopy gaps and subsequently exert a mass effect on neighbouring stands. On moist soils, we found irregular growth patterns and high rates of fungal attack, while on calcareous soils, leaf traits suggested a low nitrogen assimilation rate, limiting the production of N-containing compounds. We conclude that P. serotina fails on waterlogged and calcareous soils because it is unable to allocate sufficient energy to fruiting and/or height growth. Conversely, it succeeds on well-drained, nutrient-poor soils because of high fitness which increases its invasiveness.     

Growth and biomass turnover ofHydrocharis dubia L. cultured under different nutrient conditions

Growth of a floating-leaved plant,Hydrocharis dubia L., was examined under varying nutrient conditions between 0.3 and 30 mgN l⁻¹ total inorganic nitrogen.H. dubia plants cultured under the most nutrient-rich condition showed the highest maximum ramet density (736 m⁻²), the highest maximum biomass (80.4 g dry weight m⁻²), and the highest total net production (185 g dry weight m⁻² in 82 days). Plants under nutrient-poor conditions had a relatively large proportion of root biomass and a small proportion of leaves with a long life span. Compared with other floating-leaved and terrestrial plants, the maximum biomass ofH. dubia was relatively small. This, and the rapid biomass turnover, was related to the short life span of leaves (13.2–18.7 days) and large biomass distribution to leaves.     

The influence of harvester and Messor capensis nest-mounds on the productivity and distribution of some plant species in the southern Karoo,South Africa

Nest-mounds of the harvester ant Messor capensis occur on and around nutrient-rich patches, along minor drainage lines in nutrient-rich soils, and on the plains, generally in nutrient-poor soils. Nest-site selection is related to the presence of suitable deep soils, the presence of stones and the distance from the nearest neighbouring nest. Two plant species, Galenia fruticosa and Pteronia pallens, were significantly associated with Messor capensis nest-mounds, both in numbers of mounds occupied and in numbers of individuals. A third species, Drosanthemum montaguense, was also more common in numbers of individuals, while a fourth species, Rhinephyllum macradenium was negatively associated with these mounds. The analysis of species guilds by soil type shows that significantly more species of nutrient-rich soils are present on M. capensis nest-mounds. Also, significantly more taller, woody species occurred on nest-mounds than in inter-mound spaces. Two species, Pteronia pallens and Osteospermum sinuatum, growing on ant nest-mounds had significantly longer inter-nodes than the same species growing off mounds. However, two other species, Pteronia cf. empetrifolia and Galenia fruticosa showed no difference in inter-node lengths between plants growing on and off mounds. Six of the nine species of plants sampled on ant nest-mounds had significantly higher seed production than plants of the same species growing in inter-mound spaces. The other three species showed a tendency towards more seeds per plant on ant nest-mounds. The proportions of live and dead plants on mounds differed between species. Only Ruschia spinosa showed a significant difference between the numbers of dead plants in the population on and off mounds, with more dead plants occurring on mounds. Significantly more seeds set on individuals of Pteronia pallens growing on ant nest-mounds than those growing off nest-mounds, but no such difference occurred in P. cf. empetrifolia. There was no significant difference in the proportion of seeds parasitized by the tephritid fly Desmella anceps for individuals of P. pallens and P. cf. empetrifolia growing on and off mounds.     

Dry ash,a better reference base than dry matter for heavy metals and other persistent pollutants

Concentrations of pollutants in biotic materials are usually related to dry matter. This work demonstrates with the use of heavy metal-contaminated poplar leaves (Populus nigra ‘Italica’), that ash of dry matter (dry ash) obviously yields a better reference, as it eliminates the natural variations among organic substance contents. Values of heavy-metal concentrations in dry ash of leaves were generally more homogeneous and better correlated with corresponding values in air and soil than those in dry matter. It is further shown that concentrations of non-biodecomposable and non-volatile pollutants in the mineral fraction (the same as ash) of, for instance, plants and their residues (autumn leaves, mulch compost) can directly be compared with those in soil (nearly 100% mineral), and vice versa. This facilitates the setting of maximally permissible values for pollutant concentrations in plant dry matter or their residues, and in soil, which can be used in the protection of soils against pollutants accumulated in fresh or fallen leaves, mulch or compost. In this connection, the heavy-metal and lindane margins in West German regulations for issuing environmental certificates for compost products are discussed.     

Site fertility and soil water-table level affect fungal biomass production and community composition in boreal peatland forests

A substantial amount of below-ground carbon (C) is suggested to be associated with fungi, which may significantly affect the soil C balance in forested ecosystems. Ergosterol from in-growth mesh bags and litterbags was used to estimate fungal biomass production and community composition in drained peatland forests with differing fertility. Extramatrical mycelia (EMM) biomass production was generally higher in the nutrient-poor site, increased with deeper water table level and decreased along the length of the recovery time. EMM biomass production was of the same magnitude as in mineral-soil forests. Saprotrophic fungal biomass production was higher in the nutrient-rich site. Both ectomycorrhizal (ECM) and saprotrophic fungal community composition changed according to site fertility and water table level. ECM fungal community composition with different exploration types may explain the differences in fungal biomass production between peatland forests. Melanin-rich Hyaloscypha may indicate decreased turnover of biomass in nutrient-rich young peatland forest. Genera Lactarius and Laccaria may be important in nutrient rich and Piloderma in the nutrient-poor conditions, respectively. Furthermore, Paxillus involutus and Cortinarius sp. may be important generalists in all sites and responsible for EMM biomass production during the first summer months. Saprotrophs showed a functionally more diverse fungal community in the nutrient-rich site.     

The propagule density of Lipomyces and other yeasts in Forest Soils

Yeasts constituted a major portion of the mycoflora in a variety of forest soils. Numbers ofLipomyces were highest in the mineral soil and very low in the L horizon or upper portion of the F horizon. It is felt that the form of nitrogen in the soil plays a major role in the development of large populations ofLipomyces in soil. Yeasts (excludingLipomyces) were most numerous in the L horizon and mineral soil and least common in the F horizon. Yeast cells acounted for a smaller percentage of the total fungal propagules on hardwood litter than on conifer litter.Paper No. 3740 of the Journal Series of the North Carolina State University Agr. Exp. Sta., Raleigh, N.C. 27607.     

Nutrient availability effects on vesicular-arbuscular mycorrhizal bell pepper (Capsicum annuum) seedlings and transplants

Pepper (Capsicum annuum) seeds were sown in nutrient-poor sand or nutrient-rich peat/vermiculite amended or not amended with Glomus macrocarpum. The vesicular-arbuscular mycorrhizal (VAM) seedlings were irrigated with three levels of nutrient solution, and transplanted into four levels of P-amended soil, each of which was irrigated with two levels of nutrient solution minus P. Mycorrhizal seedlings in sand were responsive to increasing nutrient levels; in nutrient-rich peat the seedlings did not respond to additional fertilisation. The greatest seedling development accompanied by good fungus colonisation was in nutrient-poor medium irrigated with the highest nutrient solution tested (18 mM N, 1.2 mM P, and 7 mM K). Non-VAM plants almost ceased growing between the weeks 4 and 5, whereas VAM plants increased in weight by 41–188%. After transplanting, sand-grown seedlings benefited from VAM when 300 mg P/kg or more was added to the soil but peat-grown plants did not. Fruit development was delayed in all non-VAM plants compared with VAM ones.     

Effects of Ledum groenlandicum amendments on soil characteristics and black spruce seedling growth

The effects of leaves and litter of the boreal forest understory shrub, Ledum groenlandicum, on soil characteristics and black spruce (Picea mariana) seedling growth were investigated. Organic and mineral soils, not previously associated with L. groenlandicum, were amended with leaves and litter of this species. The objectives of the present study were: (i) to determine the changes in soil characteristics after amending with L. groenlandicum, (ii) to determine the quantitative variation in the concentration of water-soluble phenolic allelochemicals in mineral and organic soil layers modified by L. groenlandicum and (iii) to study the growth response of black spruce in soils treated with different L. groenlandicum amendments. The amended organic and mineral soils were analyzed for pH, organic matter, PO₄, N, Ba, Cu, Zn, Fe, Mn, Ca, Na, K, Mg, Al and total phenolics equivalence. Results indicate that organic soils amended with L. groenlandicum leaves and litter were significantly different from unamended control soil for most of the chemical characteristics, while amended mineral soil was different from that of unmodified mineral soil for PO₄, organic matter, K and total phenolics equivalence. Water-soluble phenolics from L. groenlandicum and changes in nutrient availability are plausible causes of L. groenlandicum interference with black spruce seedling growth.     

Nutrient utilization in Quercus leucotrichophora and Pinus roxburghii seedlings at five soil fertility levels

Abstract. Nutrient utilization and growth performance of Pinus roxburghii Sarg. and Quercus leucotrichophora A. Camus, the two major forest trees of the Central Himalaya Mountains between 1000 and 2000 m elevation, were analysed at different nutrient levels. The early successional P. roxburghii occurs naturally on nutrient-poor sites and the late successional Q. leucotrichophora on nutrient-rich sites. Seedlings of these two species were grown both in monoculture and mixed, in plastic bags containing 1 kg soil and representing five levels of nutrients (NPK) in a 12 : 32 : 16 ratio. The P. roxburghii seedlings showed significantly greater nutrient extraction efficiency (nutrient unit extracted by seedlings per nutrient unit in the soil) than the Q. leucotrichophora seedlings, and this difference was increased in interspecific competition, particularly at the higher nutrient levels. In interspecific competition the extraction efficiencies of P. roxburghii for N, P and K were distinctly higher than those of Q. leucotrichophora. This we consider to be the main reason for the replacement of Q. leucotrichophora by P. roxburghii, even on nutrient-rich soils, after disturbance and subsequent increase in light availability. In both the species, re-translocation of nutrients from senescing leaves declined with increasing soil fertility. For example, re-translocation for P. roxburghii was 49% N, 30% P and 32% K at the lowest soil fertility, and 20% N, 8% P and 13% K at the highest soil fertility.     

Distribution du sélénium et de seize éléments dans les différents organes deTrifolium repens

Summary Sandy clay loam soils in the La Roche-Posay area, and the different parts ofTrifolium repens growing on 9 locations, were analyzed for their contents of selenium and 16 other elements. The level of K, P, Se in the young leaves was 2 or 3 times higher than in the old organs. The roots showed the lowest contents for these three elements. By contrast, their content of Al, Fe, As were 10 times higher than in the young leaves. When the content of Se in leaves and roots was low, the contents of P, Al and Fe increased with increasing contents of Se. When the level of Se was high, the reverse was the case.      

Soil nitrogen turnover in proximal and distal stem areas of European beech trees

In European beech (Fagus sylvatica L.) forests, a large proportion of the water and ion input to the soil results from stemflow which creates a soil microsite of high element fluxes proximal to the tree trunk. The soil proximal to the stem is considered to have different rates of nitrogen turnover which might influence the estimation of N-turnover rates at the stand scale. In a previous study we reported high nitrate fluxes with seepage proximal to the stems in a forest dominated by European beech in Steigerwald, Germany. Here, we investigated the soil nitrogen turnover in the top 15 cm soil in proximal (defined as 1 m² around beech stems) and distal stem areas. Laboratory incubations and in situ sequential coring incubations were used to determine the net rates of ammonification, nitrification, and root uptake of mineral nitrogen. In the laboratory incubations higher rates of net nitrogen mineralization and nitrification were found in the forest floor proximal to the stem as compared to distal stem areas. No stem related differences were observed in case of mineral soil samples. In contrast, the in situ incubations revealed higher rates of nitrification in the mineral soil in proximal stem areas, while net nitrogen mineralization was equal in proximal and distal areas. In the in situ incubations the average ratio of nitrification/ammonification was 0.85 in proximal and 0.34 in distal stem areas. The net nitrogen mineralization was 4.4 g N m^-2 90 day^-1 in both areas. Mineralized nitrogen was almost completely taken up by tree roots with ammonium as the dominant nitrogen species. The average ratio of nitrate/ammonium uptake was 0.69 in proximal and 0.20 in distal areas. The higher water content of the soil in proximal stem areas is considered to be the major reason for the increased rates of nitrification. Different nitrogen turnover rates in proximal stem areas had no influence on the nitrogen turnover rates in soil at the stand scale. Consequently, the observed high nitrate fluxes with seepage proximal to stems are attributed to the high nitrogen input by stemflow rather than to soil nitrogen turnover. This revised version was published online in August 2006 with corrections to the Cover Date.     

Anatomical characteristics of individual roots within the fine-root architecture of Chamaecyparis obtusa (Sieb. & Zucc.) in organic and mineral soil layers

Nutrient availability and temporal variation of physical stress are usually higher in organic soil layers than in mineral soils. Individual roots within the fine-root system adjust anatomical, morphological, and turnover characteristics to soil conditions, for example nutrient availability and physical stresses. We investigated anatomical traits, including cork formation and passage and protoxylem cell numbers, in cross-sections of individual fine roots of the conifer Chamaecyparis obtusa (Siebold & Zucc.) growing under different soil conditions. The fine-root systems in different soil layers were compared by sampling ingrowth cores buried for 1 year and filled with organic and mineral soil substrates. The number of exodermal passage cells was lower in roots from organic soils than in those from mineral soils, suggesting that apical roots tend to be more stress-tolerant in the organic layer than in mineral soils. In contrast, both root tip and specific root tip density were higher in roots from organic soils than in those from mineral soil layers. The proportion of roots with two strands of protoxylem (diarch) was greater in organic (90%) than in mineral (25%) soils. Thus, although the absorptivity of individual apical roots decreases in organic layers, the absorptivity of the entire fine-root system of C. obtusa may be increased as a result of the increase in apical root density and the proportion of ephemeral roots. We found that the fine-root system had simultaneous plasticity in density, anatomy, and architecture in response to complex soil conditions.     

Phenolic content of daylight-exposed and shaded floating leaves of water lilies (Nymphaeaceae) in relation to infection by fungi

Under suboptimal growing conditions (e.g. a lack of sunshine), floating leaves of Nymphaea alba and Nuphar lutea can become heavily infected with the fungi Colletotrichum nymphaeae and Pythium F, respectively. These fungi normally act as decomposers of senescent leaves. Mature leaves of Nymphaea alba and Nuphar lutea contain high concentrations of phenolics, secondary substances known for their fungistatic properties. The production of these compounds requires energy and primary metabolites. The hypothesis that suboptimal growing conditions reduce the ability of nymphaeids to maintain a sufficiently high level of phenolics, thereby making them more vulnerable to infection by fungi, was tested. Outdoor mesocosm experiments were used to examine the response of floating leaves of Nymphaea alba and Nuphar lutea to reduced light availability. Shading significantly reduced the phenolic content of the leaves. This was accompanied by higher disease severity. The outcome of this experiment is also discussed in relation to the higher nitrogen content measured in shaded leaves. Received: 8 February 1997 / Accepted: 12 July 1997     

Towards an understanding of the Holocene distribution of Fagus sylvatica L.

Aim Understanding the driving forces and mechanisms of changes in past plant distribution and abundance will help assess the biological consequences of future climate change scenarios. The aim of this paper is to investigate whether modelled patterns of climate parameters 6000 years ago can account for the European distribution of Fagus sylvatica at that time. Consideration is also given to the role of non‐climatic parameters as driving forces of the Holocene spread and population expansion of F. sylvatica. Location Europe. Methods European distributions were simulated using a physiologically‐based bioclimatic model (STASH) driven by three different atmospheric general circulation model (AGCM) outputs for 6000 years ago. Results The three simulations generally showed F. sylvatica to have potentially been as widespread 6000 years ago as it is today, which gives a profound mismatch with pollen‐based reconstructions of the F. sylvatica distribution at that time. The results indicate that drier conditions during the growing season 6000 years ago could have caused a restriction of the range in the south. Poorer growth conditions with consequently reduced competitive ability were modelled for large parts of France. Main conclusions Consideration of the entire European range of F. sylvatica showed that no single driving force could account for the observed distributional limits 6000 years ago, or the pattern of spread during the Holocene. Climatic factors, particularly drought during the growing season, are the likely major determinants of the potential range. Climatic factors are regionally moderated by competition, disturbance effects and the intrinsically slow rate of population increase of F. sylvatica. Dynamic vegetation modelling is needed to account for potentially important competitive interactions and their relationship with changing climate. We identify uncertainties in the climate and pollen data, as well as the bioclimatic model, which suggest that the current study does not identify whether or not climate determined the distribution of F. sylvatica 6000 years ago. Pollen data are better suited for comparison with relative abundance gradients rather than absolute distributional limits. These uncertainties from a study of the past, where we have information about plant distribution and abundance, argue for extreme caution in making forecasts for the future using equilibrium models.