Phosphorus mining efficiency declines with decreasing soil P concentration and varies across crop species.

High soil P concentrations hinder ecological restoration of biological communities typical for nutrient-poor soils. Phosphorus mining, i.e., growing crops with fertilization other than P, might reduce soil P concentrations. However, crop species have different P-uptake rates and can affect subsequent P removal in crop rotation, both of which may also vary with soil P concentration. In a pot experiment with three soil-P-levels (High-P: 125-155 mg POlsen/kg; Mid-P: 51-70 mg POlsen/kg; Low-P: 6-21 mg POlsen/kg), we measured how much P was removed by five crop species (buckwheat, maize, sunflower, flax, and triticale). Total P removal decreased with soil-P-level and depended upon crop identity. Buckwheat and maize removed most P from High-P and Mid-P soils and triticale removed less P than buckwheat, maize, and sunflower at every soil-P-level. The difference in P removal between crops was, however, almost absent in Low-P soils. Absolute and relative P removal with seeds depended upon crop species and, for maize and triticale, also upon soil-P-level. None of the previously grown crop species significantly affected P removal by the follow-up crop (perennial ryegrass). We can conclude that for maximizing P removal, buckwheat or maize could be grown.

Plant movements and climate warming: intraspecific variation in growth responses to nonlocal soils.

Most range shift predictions focus on the dispersal phase of the colonization process. Because moving populations experience increasingly dissimilar nonclimatic environmental conditions as they track climate warming, it is also critical to test how individuals originating from contrasting thermal environments can establish in nonlocal sites. We assess the intraspecific variation in growth responses to nonlocal soils by planting a widespread grass of deciduous forests (Milium effusum) into an experimental common garden using combinations of seeds and soil sampled in 22 sites across its distributional range, and reflecting movement scenarios of up to 1600 km. Furthermore, to determine temperature and forest-structural effects, the plants and soils were experimentally warmed and shaded. We found significantly positive effects of the difference between the temperature of the sites of seed and soil collection on growth and seedling emergence rates. Migrant plants might thus encounter increasingly favourable soil conditions while tracking the isotherms towards currently 'colder' soils. These effects persisted under experimental warming. Rising temperatures and light availability generally enhanced plant performance. Our results suggest that abiotic and biotic soil characteristics can shape climate change-driven plant movements by affecting growth of nonlocal migrants, a mechanism which should be integrated into predictions of future range shifts.

Nitrogen saturation and net ecosystem production.

Magnani et al. found that net carbon (C) sequestration of temperate and boreal forests is clearly driven by nitrogen (N) deposition. From the positive relationship between average net ecosystem production (NEP) and wet N deposition, the authors further conclude that "no signs of N saturation were apparent" in the studied forests and that this is "casting doubts on the risk of widespread ecosystem nitrogen saturation". Nitrogen additions can clearly alter net ecosystem production, but net ecosystem production cannot be used as an indicator of N saturation.

Effects of two contrasting hemiparasitic plant species on biomass production and nitrogen availability.

Hemiparasitic plants can substantially change plant community structure; the drainage of host resources has a direct negative effect on host biomass and, as a consequence, promotes non-host biomass production (parasitism pathway); on the other hand, hemiparasitic litter inputs can enhance nutrient cycling which may have an indirect positive effect on both host and non-host biomass production (litter pathway). We evaluated the net effect of both pathways on total shoot biomass (with and without the hemiparasite) and shoot biomass of graminoids, forbs and ericaceous shrubs using a removal experiment in three sites infested with the annual Rhinanthus angustifolius, and three sites infested with the biennial Pedicularis sylvatica. We addressed the potential importance of litter effects by determination of litter quantity and quality, as well as modeling N release during decomposition. In the second year after removing the hemiparasites, total plant biomass at Rhinanthus sites was 24 % higher in weeded plots than in control plots, while weeding had no significant effect at Pedicularis sites. The increase in total biomass following Rhinanthus removal was mainly due to a higher biomass of graminoids. The amount of litter produced by Rhinanthus was only half of that produced by Pedicularis; N contents were similar. The amount of N in the litter was 9 and 30 % of the amount removed by mowing for Rhinanthus and Pedicularis sites, respectively. Within 2 months, about 45 % of the N in both hemiparasitic litter types was released by decomposition. Our results suggest that in addition to the suppression of host biomass due to parasitism, positive litter feedbacks on host and non-host biomass-via an increase in nutrient availability-also affect plant community structure. We propose that, depending on the particular hemiparasite and/or site conditions, these positive litter feedbacks on shoot biomass can compensate for the negative effect of parasitism.

The response of forest plant regeneration to temperature variation along a latitudinal gradient.

BACKGROUND AND AIMS: The response of forest herb regeneration from seed to temperature variations across latitudes was experimentally assessed in order to forecast the likely response of understorey community dynamics to climate warming. METHODS: Seeds of two characteristic forest plants (Anemone nemorosa and Milium effusum) were collected in natural populations along a latitudinal gradient from northern France to northern Sweden and exposed to three temperature regimes in growth chambers (first experiment). To test the importance of local adaptation, reciprocal transplants were also made of adult individuals that originated from the same populations in three common gardens located in southern, central and northern sites along the same gradient, and the resulting seeds were germinated (second experiment). Seedling establishment was quantified by measuring the timing and percentage of seedling emergence, and seedling biomass in both experiments. KEY RESULTS: Spring warming increased emergence rates and seedling growth in the early-flowering forb A. nemorosa. Seedlings of the summer-flowering grass M. effusum originating from northern populations responded more strongly in terms of biomass growth to temperature than southern populations. The above-ground biomass of the seedlings of both species decreased with increasing latitude of origin, irrespective of whether seeds were collected from natural populations or from the common gardens. The emergence percentage decreased with increasing home-away distance in seeds from the transplant experiment, suggesting that the maternal plants were locally adapted. CONCLUSIONS: Decreasing seedling emergence and growth were found from the centre to the northern edge of the distribution range for both species. Stronger responses to temperature variation in seedling growth of the grass M. effusum in the north may offer a way to cope with environmental change. The results further suggest that climate warming might differentially affect seedling establishment of understorey plants across their distribution range and thus alter future understorey plant dynamics.

Four decades of post-agricultural forest development have caused major redistributions of soil phosphorus fractions.

Fertilisation of agricultural land causes an accumulation of nutrients in the top soil layer, among which phosphorus (P) is particularly persistent. Changing land use from farmland to forest affects soil properties, but changes in P pools have rarely been studied despite their importance to forest ecosystem development. Here, we describe the redistributions of the P pools in a four-decadal chronosequence of post-agricultural common oak (Quercus robur L.) forests in Belgium and Denmark. The aim was to assess whether forest age causes a repartitioning of P throughout the various soil P pools (labile P, slowly cycling P and occluded P); in particular, we addressed the time-related alterations in the inorganic versus organic P fractions. In less than 40 years of oak forest development, significant redistributions have occurred between different P fractions. While both the labile and the slowly cycling inorganic P fractions significantly decreased with forest age, the organic fractions significantly increased. The labile P pool (inorganic + organic), which is considered to be the pool of P most likely to contribute to plant-available P, significantly decreased with forest age (from >20 to <10% of total P), except in the 0-5 cm of topsoil, where labile P remained persistently high. The shift from inorganic to organic P and the shifts between the different inorganic P fractions are driven by biological processes and also by physicochemical changes related to forest development. It is concluded that the organic labile P fraction, which is readily mineralisable, should be taken into account when studying the bioavailable P pool in forest ecosystems.

Plasticity in response to phosphorus and light availability in four forest herbs.

The differential ability of forest herbs to colonize secondary forests on former agricultural land is generally attributed to different rates of dispersal. After propagule arrival, however, establishing individuals still have to cope with abiotic soil legacies from former agricultural land use. We focused on the plastic responses of forest herbs to increased phosphorus availability, as phosphorus is commonly found to be persistently bioavailable in post-agricultural forest soils. In a pot experiment performed under field conditions, we applied three P levels to four forest herbs with contrasting colonization capacities: Anemone nemorosa, Primula elatior, Circaea lutetiana and Geum urbanum. To test interactions with light availability, half of the replicas were covered with shade cloths. After two growing seasons, we measured aboveground P uptake as well as vegetative and regenerative performance. We hypothesized that fast-colonizing species respond the most opportunistically to increased P availability, and that a low light availability can mask the effects of P on performance. All species showed a significant increase in P uptake in the aboveground biomass. The addition of P had a positive effect on the vegetative performances of two of the species, although this was unrelated to their colonization capacities. The regenerative performance was affected by light availability (not by P addition) and was related to the species' phenology. Forest herbs can obviously benefit from the increased availability of P in post-agricultural forests, but not all species respond in the same way. Such differential patterns of plasticity may be important in community dynamics, as they affect the interactions among species.

Tree species effect on the redistribution of soil metals.

Phytostabilization of metals using trees is often promoted although the influence of different tree species on the mobilization of metals is not yet clear. Soil and biomass were sampled 33 years after planting four tree species (Quercus robur, Fraxinus excelsior, Acer pseudoplatanus, Populus 'Robusta') in a plot experiment on dredged sediment. Poplar took up high amounts of Cd and Zn and this was associated with increased Cd and Zn concentrations in the upper soil layer. The other species contained normal concentrations of Cd, Cu, Cr, Pb and Zn in their tissues. Oak acidified the soil more than the other species and caused a decrease in the concentration of metals in the upper soil layer. The pH under poplar was lower than expected and associated with high carbon concentrations in the top soil. This might be assigned to retardation of the litter decomposition due to elevated Cd and Zn concentrations in the litter.

Spatial variability and temporal stability of throughfall deposition under beech (Fagus sylvatica L.) in relationship to canopy structure.

Although the spatial variability of throughfall (TF) in forest ecosystems can have important ecological implications, little is known about the driving factors of within-stand TF variability, particularly in deciduous forests. While the spatial variability of TF water amount and H+ deposition under a dominant beech (Fagus sylvatica L.) tree was significantly higher in the leafed period than in the leafless period, the spatial TF deposition patterns of most major ions were similar in both periods. The semiannual TF depositions of all ions other than H+ were significantly positively correlated (r=0.68-0.90, p<0.05) with canopy structure above sample locations throughout the entire year. The amounts of TF water and H+ deposition during the leafed period were negatively correlated with branch cover. We conclude that the spatial heterogeneity of ion deposition under beech was significantly affected by leaves in the growing period and by branches in non-foliated conditions.

Acidification of forested podzols in North Belgium during the period 1950-2000.

Acidification of forest soils in Europe and North America has been an important concern over the last decades. The last area-covering survey of forest soil acidification in Flanders (North Belgium) goes back to 1985 [Ronse A, De Temmerman L, Guns M, De Borger R. Evolution of acidity, organic matter content, and CEC in uncultivated soils of North Belgium during the past 25 years. Soil Sci; 146, (1988), 453-460] and highlighted a significant acidification of the upper layer (0.3-0.4 m) of forested podzols during the period 1950-1985. The present study aimed to assess (1) to what extent further acidification of forested podzols occurred during the period 1985-2000 at different depths and (2) whether the average annual acidification rate accelerated or slowed down between 1985 and 2000 compared to the period 1950-1985. Average soil pH-KCl values of podzols in northern Belgium dropped during the period 1985-2000. This decline extends to a depth of about 50 cm but was most pronounced and significant in the A horizon. In the A(0), A(1) and A(2) horizons, average pH dropped with 0.2, 0.3 and 0.1 units, and in the B(ir) and C horizons with 0.1 units. No change in average pH value occurred in the B(h) horizon. Average annual acidification rate of the A(1) horizon was significantly higher in the period 1985-2000 than in the period 1950-1985. Changes in pH occurred in the entire soil profile during the period 1950/67-1985 likely because sulphate was the major form of acid deposition before 1985. After 1985, acid sulphur deposition decreased with more than 50% in North Belgium. In contrast, ammonium deposition almost doubled between 1950 and 1980, which may explain why soil acidification between 1985 and 2000 has been restricted to the upper soil horizons.

Variation in throughfall deposition across a deciduous beech (Fagus sylvatica L.) forest edge in Flanders.

Throughfall deposition and canopy exchange of acidifying and eutrophying compounds and major base cations were studied by means of throughfall analysis in a deciduous beech (Fagus sylvatica L.) forest edge in Belgium over a period of 1 year. Throughfall fluxes of Cl(-), NH(4)(+) and Na(+) were significantly elevated at the forest edge compared to the forest interior. As no edge effect on throughfall water volume could be detected, the observed edge enhancement effects were mainly due to dry deposition and canopy exchange patterns. Indeed, there was an elevated dry deposition of Cl(-), Na(+), K(+), Ca(2+) and Mg(2+) up to 50 m from the field/forest border. Within the forest, throughfall and dry deposition of SO(4)(2-) were highly variable and no significant differences were observed between the forest edge and the forest interior. Leaching of K(+) and Ca(2+) was reduced in the forest edge up to a distance of 30 m from the border. The measured nitrogen and acidic depositions far exceeded the current Flemish critical loads with respect to the protection of biodiversity in forests, especially at the forest edge. This points to an urgent need for controlling emissions as well as the need to consider the elevated deposition load in forest edges when calculating the critical loads in forests.

Influences of evergreen gymnosperm and deciduous angiosperm tree species on the functioning of temperate and boreal forests.

It has been recognized for a long time that the overstorey composition of a forest partly determines its biological and physical-chemical functioning. Here, we review evidence of the influence of evergreen gymnosperm (EG) tree species and deciduous angiosperm (DA) tree species on the water balance, physical-chemical soil properties and biogeochemical cycling of carbon and nutrients. We used scientific publications based on experimental designs where all species grew on the same parent material and initial soil, and were similar in stage of stand development, former land use and current management. We present the current state of the art, define knowledge gaps, and briefly discuss how selection of tree species can be used to mitigate pollution or enhance accumulation of stable organic carbon in the soil. The presence of EGs generally induces a lower rate of precipitation input into the soil than DAs, resulting in drier soil conditions and lower water discharge. Soil temperature is generally not different, or slightly lower, under an EG canopy compared to a DA canopy. Chemical properties, such as soil pH, can also be significantly modified by taxonomic groups of tree species. Biomass production is usually similar or lower in DA stands than in stands of EGs. Aboveground production of dead organic matter appears to be of the same order of magnitude between tree species groups growing on the same site. Some DAs induce more rapid decomposition of litter than EGs because of the chemical properties of their tissues, higher soil moisture and favourable conditions for earthworms. Forest floors consequently tend to be thicker in EG forests compared to DA forests. Many factors, such as litter lignin content, influence litter decomposition and it is difficult to identify specific litter-quality parameters that distinguish litter decomposition rates of EGs from DAs. Although it has been suggested that DAs can result in higher accumulation of soil carbon stocks, evidence from field studies does not show any obvious trend. Further research is required to clarify if accumulation of carbon in soils (i.e. forest floor + mineral soil) is different between the two types of trees. Production of belowground dead organic matter appears to be of similar magnitude in DA and EG forests, and root decomposition rate lower under EGs than DAs. However there are some discrepancies and still are insufficient data about belowground pools and processes that require further research. Relatively larger amounts of nutrients enter the soil-plant biogeochemical cycle under the influence of EGs than DAs, but recycling of nutrients appears to be slightly enhanced by DAs. Understanding the mechanisms underlying forest ecosystem functioning is essential to predicting the consequences of the expected tree species migration under global change. This knowledge can also be used as a mitigation tool regarding carbon sequestration or management of surface waters because the type of tree species affects forest growth, carbon, water and nutrient cycling.

Light accelerates plant responses to warming.

Competition for light has profound effects on plant performance in virtually all terrestrial ecosystems. Nowhere is this more evident than in forests, where trees create environmental heterogeneity that shapes the dynamics of forest-floor communities(1-3). Observational evidence suggests that biotic responses to both anthropogenic global warming and nitrogen pollution may be attenuated by the shading effects of trees and shrubs(4-9). Here we show experimentally that tree shade is slowing down changes in below-canopy communities due to warming. We manipulated levels of photosynthetically active radiation, temperature and nitrogen, alone and in combination, in a temperate forest understorey over a 3-year period, and monitored the composition of the understorey community. Light addition, but not nitrogen enrichment, accelerated directional plant community responses to warming, increasing the dominance of warmth-preferring taxa over cold-tolerant plants (a process described as thermophilization(6,10-12)). Tall, competitive plants took greatest advantage of the combination of elevated temperature and light. Warming of the forest floor did not result in strong community thermophilization unless light was also increased. Our findings suggest that the maintenance of locally closed canopy conditions could reduce, at least temporarily, warming-induced changes in forest floor plant communities.

Metal uptake by young trees from dredged brackish sediment: limitations and possibilities for phytoextraction and phytostabilisation.

Five tree species (Acer pseudoplatanus L., Alnus glutinosa L. Gaertn., Fraxinus excelsior L., Populus alba L. and Robinia pseudoacacia L.) were planted on a mound constructed of dredged sediment. The sediment originated from a brackish river mouth and was slightly polluted with heavy metals. This preliminary study evaluated the use of trees for site reclamation by means of phytoextraction of metals or phytostabilisation. Although the brackish nature of the sediment caused slight salt damage, overall survival of the planted trees was satisfactory. Robinia and white poplar had the highest growth rates. Ash, maple and alder had the highest survival rates (>90%) but showed stunted growth. Ash, alder, maple and Robinia contained normal concentrations of Cd, Cu, Pb and Zn in their foliage. As a consequence these species reduce the risk of metal dispersal and are therefore suitable species for phytostabilisation under the given conditions. White poplar accumulated high concentrations of Cd (8.0 mg kg(-1)) and Zn (465 mg kg(-1)) in its leaves and might therefore cause a risk of Cd and Zn input into the ecosystem because of autumn litter fall. This species is thus unsuitable for phytostabilisation. Despite elevated metal concentrations in the leaves, phytoextraction of heavy metals from the soil by harvesting stem and/or leaf biomass of white poplar would not be a realistic option because it will require an excessive amount of time to be effective.

Metal and nutrient dynamics in decomposing tree litter on a metal contaminated site.

In a forest on sandy, metal polluted soil, we examined effects of six tree species on litter decomposition rates and accompanied changes in metal (Cd, Zn) and nutrient (base cations, N, C) amounts. Decomposition dynamics were studied by means of a litterbag experiment lasting for 30 months. The decomposition peak occurred within the first year for all tree species, except for aspen. During litter decomposition, high metal litter types released part of their accumulated metals, whereas low metal litter types were characterized by a metal enrichment. Base cations, N and C were released from all litter types. Metal release from contaminated litter might involve risks for metal dispersion towards the soil. On the other hand, metal enrichment of uncontaminated litter may be ecologically relevant as it can be easily transported or serve as food source.

Forest floor leachate fluxes under six different tree species on a metal contaminated site.

Trees play an important role in the biogeochemical cycling of metals, although the influence of different tree species on the mobilization of metals is not yet clear. This study examined effects of six tree species on fluxes of Cd, Zn, DOC, H(+) and base cations in forest floor leachates on a metal polluted site in Belgium. Forest floor leachates were sampled with zero-tension lysimeters in a 12-year-old post-agricultural forest on a sandy soil. The tree species included were silver birch (Betula pendula), oak (Quercus robur and Q. petraea), black locust (Robinia pseudoacacia), aspen (Populus tremula), Scots pine (Pinus sylvestris) and Douglas fir (Pseudotsuga menziesii). We show that total Cd fluxes in forest floor leachate under aspen were slightly higher than those in the other species' leachates, yet the relative differences between the species were considerably smaller when looking at dissolved Cd fluxes. The latter was probably caused by extremely low H(+) amounts leaching from aspen's forest floor. No tree species effect was found for Zn leachate fluxes. We expected higher metal leachate fluxes under aspen as its leaf litter was significantly contaminated with Cd and Zn. We propose that the low amounts of Cd and Zn leaching under aspen's forest floor were possibly caused by high activity of soil biota, for example burrowing earthworms. Furthermore, our results reveal that Scots pine and oak were characterized by high H(+) and DOC fluxes as well as low base cation fluxes in their forest floor leachates, implying that those species might enhance metal mobilization in the soil profile and thus bear a potential risk for belowground metal dispersion.

Throughfall deposition and canopy exchange processes along a vertical gradient within the canopy of beech (Fagus sylvatica L.) and Norway spruce (Picea abies (L.) Karst).

To assess the impact of air pollution on forest ecosystems, the canopy is usually considered as a constant single layer in interaction with the atmosphere and incident rain, which could influence the measurement accuracy. In this study the variation of througfall deposition and derived dry deposition and canopy exchange were studied along a vertical gradient in the canopy of one European beech (Fagus sylvatica L.) tree and two Norway spruce (Picea abies (L.) Karst) trees. Throughfall and net throughfall deposition of all ions other than H(+) increased significantly with canopy depth in the middle and lower canopy of the beech tree and in the whole canopy of the spruce trees. Moreover, throughfall and net throughfall of all ions in the spruce canopy decreased with increasing distance to the trunk. Dry deposition occurred mainly in the upper canopy and was highest during the growing season for H(+), NH(4)(+), NO(3)(-) and highest during the dormant season for Na(+), Cl(-), SO(4)(2-) (beech and spruce) and K(+), Ca(2+) and Mg(2+) (spruce only). Canopy leaching of K(+), Ca(2+) and Mg(2+) was observed at all canopy levels and was higher for the beech tree compared to the spruce trees. Canopy uptake of inorganic nitrogen and H(+) occurred mainly in the upper canopy, although significant canopy uptake was found in the middle canopy as well. Canopy exchange was always higher during the growing season compared to the dormant season. This spatial and temporal variation indicates that biogeochemical deposition models would benefit from a multilayer approach for shade-tolerant tree species such as beech and spruce.

Comparison of forest edge effects on throughfall deposition in different forest types.

This study examined the influence of distance to the forest edge, forest type, and time on Cl-, SO4(2-), NO3(-), and NH4+ throughfall deposition in forest edges. The forests were dominated by pedunculate oak, silver birch, or Corsican/Austrian pine, and were situated in two regions of Flanders (Belgium). Along transects, throughfall deposition was monitored at distances of 0-128 m from the forest edge. A repeated-measures analysis demonstrated that time, forest type, and distance to the forest edge significantly influenced throughfall deposition of the ions studied. The effect of distance to the forest edge depended significantly on forest type in the deposition of Cl-, SO4(2-), and NO3(-): the edge effect was significantly greater in pine stands than in deciduous birch and oak stands. This finding supports the possibility of converting pine plantations into oak or birch forests in order to mitigate the input of nitrogen and potentially acidifying deposition.

The effect of forest type on throughfall deposition and seepage flux: a review.

Converting deciduous forests to coniferous plantations and vice versa causes environmental changes, but till now insight into the overall effect is lacking. This review, based on 38 case studies, aims to find out how coniferous and deciduous forests differ in terms of throughfall (+stemflow) deposition and seepage flux to groundwater. From the comparison of coniferous and deciduous stands at comparable sites, it can be inferred that deciduous forests receive less N and S via throughfall (+stemflow) deposition on the forest floor. In regions with relatively low open field deposition of atmospheric N (<10 kg N ha(-1) year(-1)), lower NH(4)(+) mean throughfall (+stemflow) deposition was, however, reported under conifers compared to deciduous forest, while in regions with high atmospheric N pollution (>10 kg N ha(-1) year(-1)), the opposite could be concluded. The higher the open field deposition of NH(4)(+), the bigger the difference between the coniferous and deciduous throughfall (+stemflow) deposition. Furthermore, it can be concluded that canopy exchange of K(+), Ca(2+) and Mg(2+) is on average higher in deciduous stands. The significantly higher stand deposition flux of N and S in coniferous forests is reflected in a higher soil seepage flux of NO(3)(-), SO(4)(2-), K(+), Ca(2+), Mg(2+) and Al(III). Considering a subset of papers for which all necessary data were available, a close relationship between throughfall (+stemflow) deposition and seepage was found for N, irrespective of the forest type, while this was not the case for S. This review shows that the higher input flux of N and S in coniferous forests clearly involves a higher seepage of NO(3)(-) and SO(4)(2-) and accompanying cations K(+), Ca(2+), Mg(2+) and Al(III) into the groundwater, making this forest type more vulnerable to acidification and eutrophication compared to the deciduous forest type.