The effects of shading and N status on root proliferation in nutrient patches by the perennial grass Agropyron desertorum in the field

Competition for light can affect exploitation of spatially heterogeneous soil resources. To evaluate the influence of shoot status on root growth responses in nutrient-rich soil patches, we studied the effects of shading and whole-plant nitrogen status on root growth in N-enriched and nonenriched patches by mature Agropyron desertorum plants growing in the field with below-ground competition. Roots in enriched patches had greater length to weight ratios (specific root length, SRL), indicating increased absorptive surface areas, compared with roots in control patches. Increased SRL was due to increased production and length of higher order laterals rather than morphological changes in roots of the same branching order. Although the pattern of root growth rates in patches was the same for shaded and unshaded plants, the magnitude of this response to enriched patches was damped by shading. Root relative growth rates (RGR) in N-enriched patches were reduced by more than 50% by short-term shading treatments (60% reduction in photosynthetic flux density), while root RGR in unenriched patches was unaffected by shading. Unexpectedly, plants with higher nitrogen status had greater root RGR in enriched patches than plants that had not received nitrogen supplement, again with no detectable effect on root RGR in the unenriched patches. Therefore, while both shading and plant N status affected the ability of roots to exploit enriched patches by proliferation, there was no stimulation or suppression of root growth in the unenriched, control patches. Thus, plants already under competitive pressure above ground for light and below ground for nutrients should be less able to rapidly respond to opportunities presented in nutrient patches and pulses.     

Interactive effects of defoliation and an AM fungus on plants and soil organisms in experimental legume–grass communities

We established a 13‐week greenhouse experiment based on replicated microcosms to test whether the effects of defoliation on grassland plants and soil organisms depend on plant species composition and the presence of arbuscular mycorrhizal (AM) fungi. The experiment constituted of three treatment factors – plant species composition, inoculation of an AM fungus and defoliation – in a fully factorial design. Plant species composition had three levels: (1) Trifolium repens monoculture (T), (2) Phleum pratense monoculture (P) and (3) mixture of T. repens and P. pratense (T+P), while the AM inoculation and the defoliation treatment had two levels: (1) no inoculation of AM fungi and (2) inoculation of the AM fungus Glomus claroideum BEG31, and (1) no trimming, and (2) trimming of all plant material to 6 cm above the soil surface three times during the experiment, respectively. At the final harvest, AM colonization rate of plant roots differed between the plant species compositions, being on average 45% in T, 33% in T+P and 4% in P. Defoliation did not affect the colonization rate in T but raised the rate from 1% to 7% in P and from 20% to 45% in T+P. Shoot production and standing shoot and root biomass were 48%, 85% and 68% lower, respectively, in defoliated than in non‐defoliated systems, while the AM fungus did not affect shoot production and root mass but reduced harvested shoot mass by 8% in non‐defoliated systems. Of the plant quality attributes, defoliation enhanced the N concentration of harvested shoot biomass by 129% and 96% in P and T+P, respectively, but had no effect in T, while the C concentration of shoot biomass was on average 2.7% lower in defoliated than in non‐defoliated systems. Moreover, defoliation reduced shoot C yield (the combined C content of defoliated and harvested shoot biomass) on average by 47% across all plant species compositions and shoot N yield by 37% in T only. In contrast to defoliation, the AM fungus did not affect shoot N and C concentrations or shoot N yield, but induced 10% lower C yield in non‐defoliated systems and 17% higher C yield in defoliated T. In roots, defoliation led to 56% and 21% higher N concentration in P and T+P, respectively, and 28% higher C concentration in P, while the mycorrhizal fungus lowered root N concentration by 9.7% in defoliated systems and had no effect on root C concentrations. In the soil, the nematode community was dominated by bacterivores and the other trophic groups were found in a few microcosms only. Bacterivores were 45% more abundant in defoliated than in non‐defoliated systems, but were not affected by plant species composition or the AM fungus. Soil inorganic N concentration was significantly increased by defoliation in T+P, while the mycorrhizal fungus reduced NH₄–N concentration by 40% in T. The results show that defoliation had widespread effects in our experimental systems, and while the effects on plant growth were invariably negative and those on bacterivorous nematodes invariably positive, most effects on plant C and N content and soil inorganic N concentration varied depending on the plant species present. In contrast, the effects of defoliation did not depend on the presence of the AM fungus, which suggests that while the relative abundance of legumes and grasses is likely to have a significant role in the response of legume–grass communities to defoliation, the role of AM fungi may be less important. In line with this, the AM fungus had only a few significant effects on plant and soil attributes in our systems and each of them was modified by defoliation and/or plant species composition. This suggests that the effects of AM fungi in legume–grass communities may largely depend on the plant species present and whether the plants are grazed or not.     

Resource competition and fire‐regulated nutrient demand in carnivorous plants of wet pine savannas

Question: What are the mechanisms by which fire reduces competition for both a short‐lived and a long‐lived species in old‐growth ground‐cover plant communities of wet pine savannas (originally Pinus palustris, replaced by P. elliottii)? Location: Outer coastal plain of southeastern Mississippi, USA. Methods: I reviewed previous competition experiments and proposed a new hypothesis to explain the relationship between fire, competition, and species co‐existence in wet longleaf pine savannas. The first study is about growth and seedling emergence responses of a short‐lived carnivorous plant, Drosera capillaris, to reduction in below‐ground competition and above‐ plus below‐ground competition. The second study deals with growth and survival responses of a long‐lived perennial carnivorous plant, Sarracenia alata, to neighbour removal and prey‐exclusion to determine if a reduction in nutrient supply increased the intensity of competition in this nutrient‐poor system. Results: Fire increased seedling emergence of the short‐lived species by reducing above‐ground competition through the destruction of above‐ground parts of plants and the combustion of associated litter. Prey exclusion did not increase competitive effects of neighbours on the long‐lived species. However, because the experiment was conducted in a year without fire, shade reduced nutrient demand, which may have obviated competition for soil nutrients between Sarracenia alata and its neighbours. Conclusion: Repeated fires likely interact with interspecific differences in nutrient uptake to simultaneously reduce both above‐ground competition and competition for nutrients in old‐growth ground cover communities in pine savannas. Restoration practitioners should consider the possibility that the composition of the plant community is just as important as fire in ensuring that frequent fires maintain species diversity.     

The effect of simulated herbivory on growth and nutrient status of focal and neighbouring early successional woody plant species

Defoliation through herbivory is well known to affect target plants and their associated belowground properties, but the response of plants and their soil environment to defoliation of their neighbours is less well understood. We performed a controlled shade‐house experiment involving three plant species that colonize New Zealand floodplains during primary succession, i.e. a palatable N₂‐fixing shrub (Carmichaelia odorata), a palatable deciduous small tree (Fuchsia excorticata) and a less palatable evergreen tree (Weinmannia racemosa). All species were grown in large pots for 40 months both singly and in two species pairs, and either one or both of the species grown in pairs were clipped to simulate herbivory. Responses of growth and foliar nutrient status to clipping varied strongly among species, with Carmichaelia having the largest response and Fuchsia having the smallest. Carmichaelia also enhanced soil microbial biomass and activity, and foliar N concentrations of Weinmannia. However, this did not translate to a net positive effect; instead Carmichaelia competitively reduced growth and foliar P concentrations of both other species. Most effects of Carmichaelia on the soil microflora, and growth and nutrient status of its neighbours, disappeared when Carmichaelia was clipped. Further, the effect of clipping Carmichaelia had a stronger impact on growth, soil activity and nutrient status of the other two species than did the clipping of those species. These results contradict expectations that N₂‐fixing plants should promote growth of other species in pioneer communities or that defoliation of N₂‐fixers exacerbate positive effects; in our study, defoliation of Carmichaelia merely mitigated the negative effects that it had on other species. They also suggest that interplay of competition and differential herbivory among coexisting plants has important implications for soil microflora and processes, relative nutrient acquisition and stoichiometry of coexisting plant species, and potentially plant community development.     

Cascading effects of belowground predators on plant communities are density‐dependent

Soil food webs comprise a multitude of trophic interactions that can affect the composition and productivity of plant communities. Belowground predators feeding on microbial grazers like Collembola could decelerate nutrient mineralization by reducing microbial turnover in the soil, which in turn could negatively influence plant growth. However, empirical evidences for the ecological significance of belowground predators on nutrient cycling and plant communities are scarce. Here, we manipulated predator density (Hypoaspis aculeifer: predatory mite) with equal densities of three Collembola species as a prey in four functionally dissimilar plant communities in experimental microcosms: grass monoculture (Poa pratensis), herb monoculture (Rumex acetosa), legume monoculture (Trifolium pratense), and all three species as a mixed plant community. Density manipulation of predators allowed us to test for density‐mediated effects of belowground predators on Collembola and lower trophic groups. We hypothesized that predator density will reduce Collembola population causing a decrease in nutrient mineralization and hence detrimentally affect plant growth. First, we found a density‐dependent population change in predators, that is, an increase in low‐density treatments, but a decrease in high‐density treatments. Second, prey suppression was lower at high predator density, which caused a shift in the soil microbial community by increasing the fungal: bacterial biomass ratio, and an increase of nitrification rates, particularly in legume monocultures. Despite the increase in nutrient mineralization, legume monocultures performed worse at high predator density. Further, individual grass shoot biomass decreased in monocultures, while it increased in mixed plant communities with increasing predator density, which coincided with elevated soil N uptake by grasses. As a consequence, high predator density significantly increased plant complementarity effects indicating a decrease in interspecific plant competition. These results highlight that belowground predators can relax interspecific plant competition by increasing nutrient mineralization through their density‐dependent cascading effects on detritivore and soil microbial communities.     

Time‐dependent effects of fertilization on plant biomass in floating fens

Abstract. A cross‐over fertilization experiment was carried out in Dutch floating fens to investigate effects on biomass production in the same and the following years. In total 16 fertilizer treatments were applied, combining four treatments in 1999 with four treatments in 2000 (addition of 20 g.m^?2 N, 5 g.m^?2 P, both elements and unfertilized control). The above‐ground biomass production of vascular plants was co‐limited by N and P in both years. However, in plots that were only fertilized in 1999 the effects of individual nutrients differed between the two years: N‐fertilization slightly increased the amount of biomass produced in the same year (1999), whereas P‐fertilization did so in the following year (2000). Fertilizer applied in 1999 also influenced the effects of fertilizer applied in 2000. One year after N‐fertilization vascular plant growth was still co‐limited by N and P, but one year after P‐fertilization, vascular plant growth was only limited by N. Bryophyte biomass responded weakly to fertilization. Nutrient concentrations in plant biomass, nutrient standing crops and measurements of N and P availability in the soil indicated that one year after fertilization, the N‐fertilizer had mostly ‘disappeared’ from N‐fertilized plots, whereas the availability of P remained markedly enhanced in P‐fertilized plots. In addition, P‐fertilization enhanced the uptake of N by plants the following year. The time‐dependence of fertilizer effects was probably caused by (1) higher addition of P than of N relative to the requirements of plants; (2) longer retention of P than of N in the system; (3) positive effect of P‐fertilization on the availability of N; (4) contrasting effects of N‐ and P‐fertilization on nutrient losses by plants and/or on their responses to subsequent nutrient addition; (5) changing interactions between vascular plants and mosses (mainly Sphagnum spp.); (6) nutrient export through the repeated harvest of above‐ground biomass. To determine which nutrient limits plant growth fertilization experiments should be short, avoiding that indirect effects of a non‐limiting nutrient influence results. To indicate how changed nutrient supply will affect an ecosystem longer‐term experiments are needed, so that indirect effects have time to develop and be detected.     

Individual and combined effects of disturbance and N addition on understorey vegetation in a subarctic mountain birch forest

Questions: What are the effects of repeated disturbance and N‐fertilization on plant community structure in a mountain birch forest? What is the role of enhanced nutrient availability in recovery of understorey vegetation after repeated disturbance? How are responses of soil micro‐organisms to disturbance and N‐fertilization reflected in nutrient allocation patterns and recovery of understorey vegetation after disturbance? Location: Subarctic mountain birch forest, Finland. Methods: We conducted a fully factorial experiment with annual treatments of disturbance (two levels) and N‐fertilization (four levels) during 1998–2002. We monitored treatment effects on above‐ground plant biomass, plant community structure and plant and soil nutrient concentrations. Results: Both disturbance and N‐fertilization increased the relative biomass of graminoids. The increase of relative biomass of graminoids in the disturbance treatment was over twice that of the highest N‐fertilization level, and N‐fertilization further increased their relative biomass after disturbance. As repeated disturbance broke the dominance of evergreen dwarf shrubs, it resulted in a situation where deciduous species, graminoids and herbs dominated the plant community. Although relative biomass of deciduous dwarf shrubs declined with N‐fertilization, it did not cause a shift in plant community structure, as evergreen dwarf shrubs remained dominant. Both disturbance and N‐fertilization increased the N concentration in vascular plants, whereas microbial biomass N and C were not affected by the treatments. Concentrations of NH₄⁺, dissolved organic N (DON) and dissolved organic C (DOC) increased in the soil after N‐fertilization, whereas concentrations of NH₄⁺ and DON decreased after disturbance. Conclusions: Disturbances caused by e.g. humans or herbivores contribute more to changes in the understorey vegetation structure than increased levels of N in subarctic vegetation. Fertilization accelerated the recovery potential after repeated disturbance in graminoids. Microbial activities did not limit plant growth.     

Impacts of moso bamboo (Phyllostachys pubescens) invasion on dry matter and carbon and nitrogen stocks in a broad‐leaved secondary forest located in Kyoto,western Japan

In western and central Japan, the expansion of exotic moso bamboo (Phyllostachys pubescens Mazel ex J. Houz.) populations into neighboring vegetation has become a serious problem. Although the effects of bamboo invasion on biodiversity have been well studied, shifts in nutrient stocks and cycling, which are fundamental for ecosystem functioning, are not fully understood. To explore the effects of P. pubescens invasion on ecosystem functions we examined above‐ and below‐ground dry matter and carbon (C) and nitrogen (N) stocks in a pure broad‐leaved tree stand, a pure bamboo stand, and two tree–bamboo mixed stands with different vegetation mix ratios in the secondary forest of Kyoto, western Japan. In the process of invasion, bamboo shoots offset broad‐leaved tree deaths; thus, no clear trend was apparent in total above‐ or below‐ground biomass or in plant C and N stocks during invasion. However, the ratio of above‐ground to below‐ground biomass (T/R ratio at the stand level) decreased with increasing bamboo dominance, especially in the early stages of invasion. This shift indicates that rapid bamboo rhizomatous growth is a main driver of substantial changes in stand structure. We also detected rises in the C/N ratio of forest‐floor organic matter during bamboo invasion. Thus major impacts of P. pubescens invasion into broad‐leaved forests include not only early shifts in biomass allocation, but also changes in the distribution pattern of C and N stored in plants and soil.     

New evidence indicates the coarse soil fraction is of greater relevance to plant nutrition than previously suggested

Background and aims

Plant nutrient uptake from coarse soil (2–4 mm diameter) has been demonstrated for only a limited number of nutrients, and the nutritional contribution of coarse soil when present with fine soil (material <2 mm diameter) in realistic ratios is unknown. We conducted a seedling pot trial to investigate the functional relevance of this soil fraction to plant nutrition.

Methods

Fine soil was mixed with either coarse soil, or the equivalent volume of inert glass chips, in ratios identical to those occurring naturally in soil sampled from two depths at each of two sites. Seedlings of Nothofagus solandri var. cliffortioides and Weinmannia racemosa were planted in the soil mixtures and harvested after 9 months.

Results

The content of nitrogen, phosphorous, potassium, magnesium and other elements in the above ground seedling tissue was significantly increased by the presence of coarse soil. The coarse soil fraction also contributed proportionally much more to plant nutrient uptake than fine soil on a mass per mass basis.

Conclusions

Coarse soil is excluded from conventional soil analysis, so is possible that soil nutrient capital is systematically underestimated. This has implications for land management and studies of plant dynamics in relation to nutrient supply.     

Plant and soil microbial responses to defoliation in temperate semi-natural grassland

There is much interest in understanding the nature of feedback mechanisms between plants and soil organisms in grazed ecosystems. In this study, we examine the effects of different intensities of defoliation on the growth of three dominant grass species, and observe how these plant responses relate to the biomass and activity of the microbial community in the root zone. Our data show that grassland plants with varying tolerances to grazing have markedly different growth responses to defoliation, and that these responses vary with the intensity of cutting. Defoliation of grasses which are tolerant to grazing, namely Festuca rubra and Cynosurus cristatus, leads to a reduction in root mass and an increase in the allocation of resources to shoots. In contrast, defoliation of a grass with low tolerance to grazing, Anthoxanthum odoratum, had little effect on root mass, but increased the relative allocation of resources below-ground. In all plant species, defoliation led to an increase in soil microbial biomass and C use efficiency in the root zone. This response was greatest in the root zone of A. odoratum and is likely to be related to changes in root exudation pattern following defoliation. The significance of these changes in relation to soil nutrient dynamics and plant nutrient uptake during regrowth require further exploration. This revised version was published online in June 2006 with corrections to the Cover Date.     

Phosphorus and Defoliation Interact and Improve the Growth and Composition of the Plant Community and Soil Properties in an Alpine Pasture of Qinghai-Tibet Plateau

Pasture degradation caused by overgrazing and inappropriate fertiliser management is a major production and environmental threat in Qinghai-Tibet Plateau. Previous research has focused on the effects of mixed nitrogen (N) and phosphorus (P) fertiliser and reduced grazing pressure on the plant community of the grassland; however, the role of P and how it interacts with various defoliation (the process of the complete or partial removal of the above-ground parts of plants by grazing or cutting) intensities on the plant and soil of the grassland ecosystem have not been quantified. A field experiment was conducted to quantify how P application in combination of defoliation pressure could impact the dynamic change of the plant and soil in a native alpine grassland ecosystem of the Qinghai-Tibet Plateau, China, from May 2012 to September 2014. A split-plot design with 4 replicates and repeated measures was used to determine the growth and composition of plant community and soil physical and chemical properties under various levels of P fertiliser and defoliation intensity. The results showed that applying 20 kg P/ha increased the herbage yield of Melissitus ruthenica by 68% and total pasture yield by 25%. Close defoliation favoured the growth and plant frequency of the shorter species, whereas lax defoliation favoured that of the taller plant species. Medium P rate and cutting to 3 cm above ground gave an overall best outcome in pasture yield, quality and frequency and soil moisture and nutrient concentration. Application of P fertiliser with a moderate defoliation pressure to promote legume growth and N fixation has the potential to achieve multiple benefits in increasing pasture and livestock production and improving environmental sustainability in the alpine pasture of Qinghai-Tibet Plateau, a fragile and P-deficient ecosystem zone in China and its western neighbouring countries.     

Effects of defoliation intensity on soil food-web properties in an experimental grassland community

We established a greenhouse experiment based on replicated mini‐ecosystems to evaluate the effects of defoliation intensity on soil food‐web properties in grasslands. Plant communities, composed of white clover (Trifolium repens), perennial ryegrass (Lolium perenne) and plantain (Plantago lanceolata) with well‐established root and shoot systems, were subjected to five defoliation intensity treatments: no trimming (defoliation intensity 0, or DI 0), and trimming of all plant material to 35 cm (DI 1), 25 cm (DI 2), 15 cm (DI 3) and 10 cm (DI 4) above soil surface every second week for 14 weeks. Intensification of defoliation reduced shoot production and standing shoot and root mass of plant communities but increased their root to shoot ratio. Soil microbial activity and biomass decreased with intensification of defoliation. Concentrations of NO₃–N in soil steadily increased with intensifying defoliation, whereas NH₄–N concentrations did not vary between treatments. Numbers of microbi‐detritivorous enchytraeids, bacterial‐feeding rotifers and bacterial‐feeding nematodes steadily increased with intensifying defoliation, while the abundance of fungal‐feeding nematodes was significantly enhanced only in DI 3 and DI 4 relative to DI 0. The abundance of herbivorous nematodes per unit soil mass was lower in DI 3 and DI 4 than in DI 0, DI 1 and DI 2, but when calculated per unit root mass, their abundance tended to increase with defoliation intensity. The abundance of omnivorous and predatory nematodes appeared to be highest in the most intensely defoliated systems. The ratio of abundance of fungal‐feeding nematodes to that of bacterial‐feeding nematodes was not significantly affected by defoliation intensity. The results infer that defoliation intensity may significantly alter the structure of soil food webs in grasslands, and that defoliation per se is able to induce patterns observed in grazing studies in the field. The results did not support hypotheses that defoliation per se would cause a shift between the bacterial‐based and fungal‐based energy channels in the decomposer food web, or that herbivore and detritivore densities in soil would be highest under intermediate defoliation. Furthermore, our data for microbes and microbial feeders implies that the effects of defoliation intensity on soil food‐web structure may depend on the duration of defoliation and are therefore likely to be dynamic rather than constant in nature.     

Ecological and site historical aspects of N dynamics and current N status in temperate forests

Ecological developments during Holocene age and high atmospheric depositions since industrialization have changed the N dynamics of temperate forest ecosystems. A number of different parameters are used to indicate whether the forests are N‐saturated or not, most common among them is the occurrence of nitrates in the seepage water below the rooting zone. The use of different definitions to describe N saturation implies that the N status of ecosystems is not always appropriately assessed. Data on N dynamics from 53 different German forests were used to classify various development states of forest ecosystems according to the forest ecosystem theory proposed by Ulrich for which N balances of input – (output plus plant N increment) were used. Those systems where N output equals N input minus plant N increment are described as (quasi‐) Steady State Type. Those forests where N output does not equal N input minus plant N increment as in a ‘transient state.’ Forests of the transient state may lose nitrogen from the soil (Degradation Type) or gain nitrogen [e.g., from atmospheric depositions (Accumulation Type)]. Forest ecosystems may occur in four different N states: (a) (quasi‐) Steady State Type with mull type humus, (b) Degradation Type with mull type humus, (c) Accumulation Type with moder type humus, and (d) (quasi‐) Steady State Type with moder type humus. Forests with the (quasi‐) steady state with mull type humus in the forest floor (n= 8) have high‐soil pH values, high N retention by plant increment, high N contents in the mineral soils, and have not undergone large changes in the N status. Forests of the Degradation Type lose nitrogen from the mineral soil (currently degradation is occurring on one site). Most forests that have moder or mor type humus and low‐soil pH values, and low N contents in the mineral soil have gone through the transient state of organic matter loss in the mineral soils. They accumulate organic matter in the forest floor (accumulation phase, currently 21 sites are accumulating 6–21 kg N ha^?1 yr^?1) or have reached a new (quasi‐) steady state with moder/mor type humus (n= 15). N retention in the accumulation phase has significantly increased in soil with N deposition (r²= 0.38), soil acidity (considering thickness of the forest floor as indices of soil acidity, r²= 0.43) and acid deposition (sulfate deposition, r²= 0.39). Retention of N (4–20 kg N ha^?1 yr^?1) by trees decreased and of soils increased with a decrease in the availability of base cations indicating the important role of trees for N retention in less acid soils and those of soils in more acid soils. Ecosystem theory could be successfully applied on the current data to understand the dynamics of N in temperate forest ecosystems.     

The effect of reindeer grazing on decomposition, mineralization and soil biota in a dry oligotrophic Scots pine forest

Reindeer grazing in the Fennoscandian area has a considerable influence on the ground vegetation, and this is likely in turn to have important consequences for the soil biota and decomposition processes. The effects of reindeer grazing on soil biota, decomposition and mineralization processes, and ecosystem properties in a lichen‐dominated forest in Finnish Lapland were studied inside and outside a large long term fenced reindeer exclosure area. Decomposition rates of Vaccinium myrtillus leaves in litter bags were retarded in the grazed area relative to the ungrazed area, as well as in subplots from which lichens had been artificially removed to simulate grazing. The effect of reindeer grazing on soil respiration and microbial C was positive in the lichen and litter layers of the soil profile, but retarded in the humus layer. There was no effect of grazing on gross N mineralization and microbial biomass N in the humus and upper mineral soil layer, but net N mineralization was increased by grazing. In these layers soil respiration was reduced by grazing, indicating that reindeer effects reduce the ratio of C to N mineralized by soil microorganisms. Grazing stimulated populations of all trophic groupings of nematodes in the lichen layer and microbe feeding nematodes in the litter layer, indicating that grazing by reindeer has multitrophic effects on the decomposer food‐web. Grazing decreased lichen and dwarf shrub biomasses and increased the mass of litter present in the litter layer on an areal basis, but did not significantly alter total C storage per unit area in the humus and mineral soil layers. The N concentration of lichens was increased by grazing, but the N concentrations of both living and dead Pinus sylvestris needles and Empetrum hermaphroditum leaves were not affected.
There was some evidence for each of three mechanisms which could account for the grazing effects that we observed in our study. Firstly, reindeer may have changed the composition and quality of litter input by affecting plant species composition and through addition of N from urine and faeces, resulting in a lack of available C relative to N for decomposer organisms. Secondly, the organic matter in the soil may be older in the grazed area, because of reduction of recent production of lichen litter relative to the ungrazed area. The organic matter in the grazed area may have been in a different phase of decomposition from that in the exclosure. Thirdly, the soil microclimate is likely to be affected by reindeer grazing through physical removal of lichen cover on the ground, and this can have a significant influence on soil microbial processes. This is supported by the strong observed effects of experimental removal of lichens on decomposer processes. The impact of reindeer grazing on soil processes may be a result of complex interactions between different mechanisms, and this could help to explain why the below‐ground effects of reindeer grazing have different consequences to those which have been observed in recent investigations on other grazing systems.     

The Microbial Efficiency‐Matrix Stabilization (MEMS) framework integrates plant litter decomposition with soil organic matter stabilization: do labile plant inputs form stable soil organic matter?

The decomposition and transformation of above‐ and below‐ground plant detritus (litter) is the main process by which soil organic matter (SOM) is formed. Yet, research on litter decay and SOM formation has been largely uncoupled, failing to provide an effective nexus between these two fundamental processes for carbon (C) and nitrogen (N) cycling and storage. We present the current understanding of the importance of microbial substrate use efficiency and C and N allocation in controlling the proportion of plant‐derived C and N that is incorporated into SOM, and of soil matrix interactions in controlling SOM stabilization. We synthesize this understanding into the Microbial Efficiency‐Matrix Stabilization (MEMS) framework. This framework leads to the hypothesis that labile plant constituents are the dominant source of microbial products, relative to input rates, because they are utilized more efficiently by microbes. These microbial products of decomposition would thus become the main precursors of stable SOM by promoting aggregation and through strong chemical bonding to the mineral soil matrix.     

Tree physiological responses to above-ground herbivory directly modify below-ground processes of soil carbon and nitrogen cycling

Above‐ground herbivory is ubiquitous in terrestrial ecosystems, yet its impacts on below‐ground processes and consequences for plants remain ambiguous. To examine whether physiological responses of individual trees may potentially modify soil nutrient availability, we subjected Fagus sylvatica L. (European beech) and Abies alba Mill. (silver fir) to simulated foliar herbivory over two growing seasons. Above‐ground herbivory enhanced N mineralization and inorganic N availability in the soil. The total input of C from the plant roots to the soil is not known; however, carbon sequestration in the soil, measured using stable isotopic techniques, was unaffected by herbivory. Fagus responded to herbivory by producing larger leaves, with increased photosynthetic capacity and N content, which largely compensated for the loss of biomass; Abies exhibited no such response. We conclude that despite large interspecific differences in the growth response, tree physiological responses to foliar herbivory are capable of directly modifying soil biological processes.     

Identifying qualitative effects of different grazing types on below‐ground communities and function in a long‐term field experiment

Herbivory is an important modulator of plant biodiversity and productivity in grasslands, but our understanding of herbivore‐induced changes on below‐ground processes and communities is limited. Using a long‐term (17 years) experimental site, we evaluated impacts of rabbit and invertebrate grazers on some soil functions involved in carbon cycling, microbial diversity, structure and functional composition. Both rabbit and invertebrate grazing impacted soil functions and microbial community structure. All functional community measures (functions, biogeochemical cycling genes, network association between different taxa) were more strongly affected by invertebrate grazers than rabbits. Furthermore, our results suggest that exclusion of invertebrate grazers decreases both microbial biomass and abundance of genes associated with key biogeochemical cycles, and could thus have long‐term consequences for ecosystem functions. The mechanism behind these impacts are likely to be driven by both direct effects of grazing altering the pattern of nutrient inputs and by indirect effects through changes in plant species composition. However, we could not entirely discount that the pesticide used to exclude invertebrates may have affected some microbial community measures. Nevertheless, our work illustrates that human activity that affects grazing intensity may affect ecosystem functioning and sustainability, as regulated by multi‐trophic interactions between above‐ and below‐ground communities.     

Defoliation reduces soil biota – and modifies stimulating effects of elevated CO2

To understand the responses to external disturbance such as defoliation and possible feedback mechanisms at global change in terrestrial ecosystems, it is necessary to examine the extent and nature of effects on aboveground–belowground interactions. We studied a temperate heathland system subjected to experimental climate and atmospheric factors based on prognoses for year 2075 and further exposed to defoliation. By defoliating plants, we were able to study how global change modifies the interactions of the plant–soil system. Shoot production, root biomass, microbial biomass, and nematode abundance were assessed in the rhizosphere of manually defoliated patches of Deschampsia flexuosa in June in a full‐factorial FACE experiment with the treatments: increased atmospheric CO₂, increased nighttime temperatures, summer droughts, and all of their combinations. We found a negative effect of defoliation on microbial biomass that was not apparently affected by global change. The negative effect of defoliation cascades through to soil nematodes as dependent on CO₂ and drought. At ambient CO₂, drought and defoliation each reduced nematodes. In contrast, at elevated CO₂, a combination of drought and defoliation was needed to reduce nematodes. We found positive effects of CO₂ on root density and microbial biomass. Defoliation affected soil biota negatively, whereas elevated CO₂ stimulated the plant–soil system. This effect seen in June is contrasted by the effects seen in September at the same site. Late season defoliation increased activity and biomass of soil biota and more so at elevated CO₂. Based on soil biota responses, plants defoliated in active growth therefore conserve resources, whereas defoliation after termination of growth results in release of resources. This result challenges the idea that plants via exudation of organic carbon stimulate their rhizosphere biota when in apparent need of nutrients for growth.     

Defoliation of a grass is mediated by the positive effect of dung deposition,moss removal and enhanced soil nutrient contents: results from a reindeer grazing simulation experiment

Herbivory is one of the key drivers shaping plant community dynamics. Herbivores can strongly influence plant productivity directly through defoliation and the return of nutrients in the form of dung and urine, but also indirectly by reducing the abundance of neighbouring plants and inducing changes in soil processes. However, the relative importance of these processes is poorly understood. We, therefore, established a common garden experiment to study plant responses to defoliation, dung addition, moss cover, and the soil legacy of reindeer grazing. We used an arctic tundra grazed by reindeer as our study system, and Festuca ovina, a common grazing‐tolerant grass species as the model species. The soil legacy of reindeer grazing had the strongest effect on plants, and resulted in higher growth in soils originating from previously heavily‐grazed sites. Defoliation also had a strong effect and reduced shoot and root growth and nutrient uptake. Plants did not fully compensate for the tissue lost due to defoliation, even when nutrient availability was high. In contrast, defoliation enhanced plant nitrogen concentrations. Dung addition increased plant production, nitrogen concentrations and nutrient uptake, although the effect was fairly small. Mosses also had a positive effect on aboveground plant production as long as the plants were not defoliated. The presence of a thick moss layer reduced plant growth following defoliation. This study demonstrates that grasses, even though they suffer from defoliation, can tolerate high densities of herbivores when all aspects of herbivores on ecosystems are taken into account. Our results further show that the positive effect of herbivores on plant growth via changes in soil properties is essential for plants to cope with a high grazing pressure. The strong effect of the soil legacy of reindeer grazing reveals that herbivores can have long‐lasting effects on plant productivity and ecosystem functioning after grazing has ceased.     

Effects of a soil fulvic acid on the growth and nutrient content of cucumber (Cucumis sativus) plants

Summary Cucumber (Cucumis sativus) plants were grown in Hoagland solution to which 20 to 2000 ppm of a soil fulvic acid (FA) were added. The addition of 100 to 300 ppm of FA produced highly significant increases in the growth and development of above and below ground plant parts, in the uptake of nutrient elements (N, P, K, Ca, Mg, Cu, Fe and Zn), and in the formation of numbers of flowers per plant. Effects of adding 500 and more ppm of FA were less beneficial.