Mycorrhizosphere responsiveness to atmospheric ozone and inoculation with Phytophthora citricola in a phytotron experiment with spruce/beech mixed cultures

The aim was to analyze functional changes in the mycorrhizosphere (MR) of juvenile spruce and beech grown in a mixture under ambient and twice ambient ozone and inoculated with the root pathogen Phytophthora citricola. The phytotron experiment was performed over two vegetation periods, adding the pathogen at the end of the first growing season. Root biomass data suggest that the combined treatment affected spruce more than beech and that the negative influence of ozone on stress tolerance against the root pathogen P. citricola was greater for spruce than for beech. In contrast, beech was more affected when the pathogen was the sole stressor. The functional soil parameter chosen for studies of MR soil samples was activity of extracellular enzymes. After the first year of ozone exposure, MR soil samples of both species showed increased activity of almost all measured enzymes (acid phosphatase, chitinase, beta-glucosidase, cellobiohydrolase) in the O3 treatment. Species-specific differences were observed, with a stronger effect of P. citricola on beech MR and a stronger ozone effect on spruce MR. In the second year, the effects of the combined treatment (ozone and P. citricola) were a significant increase in the activity of most enzymes (except cellobiohydrolase) for both tree species. The results indicated that responsiveness of MR soils towards ozone and P. citricola was related to the severity of infection of the plants and the reduction of belowground biomass, suggesting a strong, direct influence of plant stress on MR soil enzyme activity. Additional research is needed using different species and combined stresses to determine the broader ecological relevance of shifts in rhizosphere enzymes.     

Analysis of competition effects in mono- and mixed cultures of juvenile beech and spruce by means of the plant growth simulation model PLATHO

Inter- and intra-specific competition between plants for external resources is a critical process for plant growth in natural and managed ecosystems. We present a new approach to simulate competition for the resources light, water, and nitrogen between individual plants within a canopy. This approach was incorporated in a process-oriented plant growth simulation model. The concept of modelling competition is based on competition coefficients calculated from the overlap of occupied crown and soil volumes of each plant individual with the occupied volumes of its four nearest neighbours. The model was parameterised with data from a two-year phytotron experiment with juvenile beech and spruce trees growing in mono- and mixed cultures. For testing the model, an independent data set from this experiment and data from a second phytotron experiment with mixed cultures were used. The model was applied to analyse the consequences of start conditions and plant density on plant-plant competition. In both experiments, spruce dominated beech in mixed cultures. Based on model simulations, we postulate a large influence of start conditions and stand density on the outcome of the competition between the species. When both species have similar heights at the time of canopy closure, the model suggests a greater morphological plasticity of beech compared with spruce to be the crucial mechanism for competitiveness in mixed canopies. Similar to the experiment, in the model greater plasticity was a disadvantage for beech leading to it being outcompeted by the more persistent spruce.     

Short term effects of ozone on the plant-rhizosphere-bulk soil system of young beech trees

Plant growth largely depends on microbial community structure and function in the rhizosphere. In turn, microbial communities in the rhizosphere rely on carbohydrates provided by the host plant. This paper presents the first study on ozone effects in the plant-rhizosphere-bulk soil system of 4-year-old beech trees using outdoor lysimeters as a research platform. The lysimeters were filled with homogenized soil from the corresponding horizons of a forest site, thus minimizing field heterogeneity. Four lysimeters were treated with ambient ozone (1 x O3) and four with double ambient ozone concentrations (2 x O3; restricted to 150 ppb). In contrast to senescence, which was almost unaffected by ozone treatment, both the photochemical quantum yield of photosystem II (PSII) and leaf gas exchange were reduced (11 - 45 %) under the elevated O3 regime. However, due to large variation between the plants, no statistically significant O3 effect was found. Even though the amount of primary metabolites, such as sugar and starch, was not influenced by elevated O3 concentrations, the reduced photosynthetic performance was reflected in leaf biochemistry in the form of a reduction in soluble phenolic metabolites. The rhizosphere microbial community also responded to the O3 treatment. Both community structure and function were affected, with a tendency towards a lower diversity and a significant reduction in the potential nutrient turnover. In contrast, litter degradation was unaffected by the fumigation, indicating that in situ microbial functionality of the bulk soil did not change.     

Enhanced ozone exposure of European beech (Fagus sylvatica) stimulates nitrogen mobilization from leaf litter and nitrogen accumulation in the soil

Abstract

In a lysimeter study with young beech trees, the effects of elevated ozone concentration on the decomposition and fate of nitrogen in ¹⁵N‐labeled leaf litter were analyzed after one growing season. Nitrogen in the litter was dominated by a relatively inert, residual fraction, but easily decomposable nitrogen was present in substantial amounts. Nitrogen loss was significantly higher at twice‐ambient ozone which was largely attributed to an enhanced mobilization of residual nitrogen. Enhanced mobilization of nitrogen from litter at twice‐ambient ozone exposure resulted in additional ¹⁵N incorporation into the soil down to 30 cm depth. Only 0.41–0.62% of the nitrogen in the litter was incorporated into plant material at both ozone concentrations. Twice‐ambient ozone exposure changed the distribution of the nitrogen taken up from litter inside the beech trees in favor of the shoot, where it may have been used in biosynthetic processes required for defense reactions.     

A dynamical model of environmental effects on allocation to carbon-based secondary compounds in juvenile trees

BACKGROUND AND AIMS: Patterns and variations in concentration of carbon-based secondary compounds in plant tissues have been explained by means of different complementary and, in some cases, contradictory plant defence hypotheses for more than 20 years. These hypotheses are conceptual models which consider environmental impacts on plant internal demands. In the present study, a mathematical model is presented, which converts and integrates the concepts of the 'Growth-Differentiation Balance' hypothesis and the 'Protein Competition' model into a dynamic plant growth model, that was tested with concentration data of polyphenols in leaves of juvenile apple, beech and spruce trees. The modelling approach is part of the plant growth model PLATHO that considers simultaneously different environmental impacts on the most important physiological processes of plants. METHODS: The modelling approach for plant internal resource allocation is based on a priority scheme assuming that growth processes have priority over allocation to secondary compounds and that growth-related metabolism is more strongly affected by nitrogen deficiency than defence-related secondary metabolism. KEY RESULTS: It is shown that the model can reproduce the effect of nitrogen fertilization on allocation patterns in apple trees and the effects of elevated CO(2) and competition in juvenile beech and spruce trees. The analysis of model behaviour reveals that large fluctuations in plant internal availability of carbon and nitrogen are possible within a single vegetation period. Furthermore, the model displays a non-linear allocation behaviour to carbon-based secondary compounds. CONCLUSIONS: The simulation results corroborate the underlying assumptions of the presented modelling approach for resource partitioning between growth-related primary metabolism and defence-related secondary metabolism. Thus, the dynamical modelling approach, which considers variable source and sink strengths of plant internal resources within different phenological growth stages, presents a successful translation of existing concepts into a dynamic mathematical model.     

Acclimation to ozone affects host/pathogen interaction and competitiveness for nitrogen in juvenile Fagus sylvatica and Picea abies trees infected with Phytophthora citricola

In a two-year phytotron study, juvenile trees of European beech (Fagus sylvatica) and Norway spruce (Picea abies) were grown in mixture under ambient and twice ambient ozone (O3) and infected with the root pathogen Phytophthora citricola. We investigated the influence of O3 on the trees' susceptibility to the root pathogen and assessed, through a 15N-labelling experiment, the impact of both treatments (O3 exposure and infection) on belowground competitiveness. The hypotheses tested were that: (1) both P. citricola and O3 reduce the belowground competitiveness (in view of N acquisition), and (2) that susceptibility to P. citricola infection is reduced through acclimation to enhanced O3 exposure. Belowground competitiveness was quantified via cost/benefit relationships, i.e., the ratio of structural investment in roots relative to their uptake of 15N. Beech had a lower biomass acquisition and captured less 15N under enhanced O3 and P. citricola infection alone than spruce, whereas the latter species appeared to profit from the lower resource acquisition of beech in these treatments. Nevertheless, in the combined treatment, susceptibility to P. citricola of spruce was increased, while beech growth and 15N uptake were not further reduced below the levels found under the single treatments. Potential trade-offs between stress defence, growth performance, and associated nitrogen status are discussed for trees affected through O3 and/or pathogen infection. With respect to growth performance, it is concluded that O3 enhances susceptibility to the pathogen in spruce, but apparently raises the defence capacity in beech..     

生活污水慢渗生态处理对土壤及杨树生长的影响

为了观测生活污水杨树林地处理对土壤和林木生长的影响,2008—2009年在郑州市龙湖镇,采用不同水力负荷(0、3、6、9、12、15cm/周),进行了污水慢渗生态处理试验。测定了污水处理期间杨树地上部分生长量,对表层(0—40cm)和下层(40—100cm)土壤理化性质进行了分析。选用土壤容重、有机质、全氮、全磷、全钾、速效氮、速效磷、速效钾作为土壤质量因子,运用土壤质量综合指标评价不同处理表层土壤质量,对下层土壤的理化性状的变化进行了研究。结果表明:在3—9cm/周水力负荷时,表层土壤质量综合指标值(QI值)和杨树地上部分生长量均随着水力负荷的增加而增加,在9cm/周水力负荷时达到最大;水力负荷大于9cm/周,QI值和杨树地上部分生长量随着水力负荷增加而降低。在水力负荷较低时,污水处理对下层土壤性状影响较小,水力负荷大于9cm/周,污水处理对下层土壤性状产生了不良影响。生活污水杨树林地处理比较适宜的水力负荷是6—9cm/周。     

CASIROZ: Root parameters and types of ectomycorrhiza of young beech plants exposed to different ozone and light regimes

Tropospheric ozone (O(3)) triggers physiological changes in leaves that affect carbon source strength leading to decreased carbon allocation below-ground, thus affecting roots and root symbionts. The effects of O(3) depend on the maturity-related physiological state of the plant, therefore adult and young forest trees might react differently. To test the applicability of young beech plants for studying the effects of O(3) on forest trees and forest stands, beech seedlings were planted in containers and exposed for two years in the Kranzberg forest FACOS experiment (Free-Air Canopy O(3) Exposure System, http://www.casiroz.de ) to enhanced ozone concentration regime (ambient [control] and double ambient concentration, not exceeding 150 ppb) under different light conditions (sun and shade). After two growing seasons the biomass of the above- and below-ground parts, beech roots (using WinRhizo programme), anatomical and molecular (ITS-RFLP and sequencing) identification of ectomycorrhizal types and nutrient concentrations were assessed. The mycorrhization of beech seedlings was very low ( CA. 5 % in shade, 10 % in sun-grown plants), no trends were observed in mycorrhization (%) due to ozone treatment. The number of Cenococcum geophilum type of ectomycorrhiza, as an indicator of stress in the forest stands, was not significantly different under different ozone treatments. It was predominantly occurring in sun-exposed plants, while its majority share was replaced by Genea hispidula in shade-grown plants. Different light regimes significantly influenced all parameters except shoot/root ratio and number of ectomycorrhizal types. In the ozone fumigated plants the number of types, number of root tips per length of 1 to 2 mm root diameter, root length density per volume of soil and concentration of Mg were significantly lower than in control plants. Trends to a decrease were found in root, shoot, leaf, and total dry weights, total number of root tips, number of vital mycorrhizal root tips, fine root (mass) density, root tip density per surface, root area index, concentration of Zn, and Ca/Al ratio. Due to the general reduction in root growth indices and nutrient cycling in ozone-fumigated plants, alterations in soil carbon pools could be predicted.     

Competition increasingly dominates the responsiveness of juvenile beech and spruce to elevated CO2 and/or O3 concentrations throughout two subsequent growing seasons

Saplings of Fagus sylvatica and Picea abies were grown in mono‐ and mixed cultures in a 2‐year phytotron study under all four combinations of ambient and elevated ozone (O₃) and carbon dioxide (CO₂) concentrations. The hypotheses tested were (1) that the competitiveness of beech rather than spruce is negatively affected by the exposure to enhanced O₃ concentrations, (2) spruce benefits from the increase of resource availability (elevated CO₂) in the mixed culture and (3) that the responsiveness of plants to CO₂ and O₃ depends on the type of competition (i.e. intra vs. interspecific). Beech displayed a competitive disadvantage when growing in mixture with spruce: after two growing seasons under interspecific competition, beech showed significant reductions in leaf gas exchange, biomass development and crown volume as compared with beech plants growing in monoculture. In competition with spruce, beech appeared to be nitrogen (N)‐limited, whereas spruce tended to benefit in terms of its plant N status. The responsiveness of the juvenile trees to the atmospheric treatments differed between species and was dominated by the type of competition: spruce growth benefited from elevated CO₂ concentrations, while beech growth suffered from the enhanced O₃ regime. In general, interspecific competition enhanced these atmospheric treatment effects, supporting our hypotheses. Significant differences in root : shoot biomass ratio between the type of competition under both elevated O₃ and CO₂ were not caused by readjustments of biomass partitioning, but were dependent on tree size. Our study stresses that competition is an important factor driving plant development, and suggests that the knowledge about responses of plants to elevated CO₂ and/or O₃, acquired from plants growing in monoculture, may not be transferred to plants grown under interspecific competition as typically found in the field.     

Willow growth in response to nutrients and moisture on a clay landfill cap soil. I. Growth and biomass production

The growth and biomass production by willow (Salix viminalis L.) was studied in lysimeters containing Oxford clay landfill cap soil with different amendments, bulk densities and watering regimes. Three years from planting, stem biomass in well-watered plants was least (0.28 kg plant(-1)) with high bulk density soil (1480 kg m(-3)) and no nutritional amendment but was increased 10-fold (2.53 kg plant(-1)) by reducing soil bulk density (1200 kg m3) and adding amendments. In comparison, on a sandy loam soil it was 6.23 kg plant(-1). There were similar differences in number of stems plant(-1), stem basal area plant(-1) and plant leaf area which can be attributed to low nitrogen and phosphorus levels in Oxford clay. Water stress reduced stem biomass production by 26-37% and caused higher root:stem ratios. These were also higher on Oxford clay than on the sandy loam. Successful biomass production from willow on Oxford clay landfill caps will therefore require nutritional amendment.     

O3浓度升高对小麦根际土壤酶活性和有机酸含量的影响

近地层臭氧(O₃)浓度升高作为全球气候变化的重要因素之一,对土壤生态环境和农作物生长发育造成了很大影响.本研究采用开顶式气室(OTCs)法,探究臭氧浓度升高对小麦不同生育期(分蘖期、拔节期、孕穗期和成熟期)根际土壤酶活性(过氧化氢酶、多酚氧化酶、脱氢酶和转化酶)和有机酸含量(草酸、柠檬酸和苹果酸)的影响规律,并结合根际土壤理化性质、植株根系生长状况等分析其产生影响的原因.结果表明: O₃浓度升高不同程度地提高了小麦成熟期土壤过氧化氢酶、多酚氧化酶、脱氢酶和转化酶活性,其中过氧化氢酶和多酚氧化酶活性提高达显著水平;在抽穗期,脱氢酶和转化酶活性因臭氧浓度升高而显著提高,增幅最高可达76.7%.在成熟期,O₃浓度升高显著提高了根际土壤中柠檬酸和苹果酸含量;显著降低了根际土壤pH、电导率、总碳和总氮含量,增加了土壤氧化还原电位(Eh);显著降低了小麦根系生物量、总根长和根总表面积,而增加了根平均直径.     

Influence of chronic ozone stress on carbon translocation pattern into rhizosphere microbial communities of beech trees (Fagus sylvatica L.) during a growing season

The influence of long-term chronic ozone exposure on carbon fluxes from young beech trees (Fagus sylvatica L.) into the phospholipid fraction of microbial communities (PLFA) in the rhizosphere and into the dissolved organic carbon (DOC) fraction was studied in a lysimeter experiment using ¹³C depleted CO₂ over one vegetation period to identify possible changes in below ground carbon translocation processes due to the plant stress. It could be shown that microbial biomass as well as individual microbial communities and their activity pattern in the rhizosphere of young beech trees are mainly driven by the vegetation period. An increase in total microbial biomass as well as individual microbial communities was detected during the vegetation period from June to September. However, also a clear ozone effect was visible mainly at the end of the vegetation period. Enzyme activities and PLFA data indicated earlier induced plant senescence as a response to the elevated ozone treatment. Furthermore higher microbial biomass and abundance of plant C utilizing microbes was observed in elevated ozone treatments over the whole vegetation period.     

Structure and activities of ectomycorrhizal and microbial communities in the rhizosphere of Fagus sylvatica under ozone and pathogen stress in a lysimeter study

The aim was to study the influence of abiotic (elevated ozone) or biotic stress (Phytophthora citricola) or their combination on soil biological components and processes in the rhizosphere of young beech trees. Ectomycorrhizal and overall microbial community composition was studied at two soil depths in a lysimeter experiment with 7 year old trees of Fagus sylvatica. As a functional parameter, potential enzyme activities were measured in mycorrhizosphere soil and on excised mycorrhizal tips. The degree of mycorrhization, structure and potential enzymatic activities of mycorrhizal communities were only slightly influenced by treatments. Soil enzyme activities were depressed under elevated ozone and stimulated by P. citricola under ambient but not under elevated ozone. Overall microbial community composition (PLFA) and ectomycorrhizal diversity changed with depth. PLFA analyses not only suggested a reaction of the microbial community to elevated ozone but also indicated an increase in plant stress related components. No influence of the biotic stress on ectomycorrhizal or overall microbial community structure was detected. Changes in the mycorrhizosphere community structure and function due to ozone may be explained by the quality of plant derived carbon.     

Ozone effects on visible foliar injury and growth of Fagus sylvatica and Viburnum lantana seedlings grown in monoculture or in mixture

Seedlings of Fagus sylvatica (beech) and Viburnum lantana (Viburnum) grown in monoculture and mixture were exposed to ambient and sub-ambient (charcoal-filtered) ozone concentrations in open-top chambers over the course of the 2003 and 2004 growing seasons at the WSL Lattecaldo open-top chamber facility in southern Switzerland. The aim of the study was to determine how the sensitivity to ozone in ambient air of these two species would differ between monocultures and mixtures in terms of growth and visible foliar injury development. Ambient ozone concentrations were consistently higher from the end of April to the middle of October in 2003 than in 2004 with seasonal peaks and means reaching 147 and 50 parts-per-billion (ppb) in 2003 compared to 124 and 40 ppb in 2004. Ambient AOT40 (ozone concentration accumulated over a threshold of 40 ppb during daylight hours with global radiation >50 W m^?2) values from the end of April to the middle of October reached 48.3 and 26.8 parts-per-billon hours (ppm h) in 2003 and 2004, respectively. In general, Viburnum was a stronger competitor than beech over the course of this 2-year study. Seedlings of Viburnum benefited from interspecific competition in terms of both height growth and above-ground biomass accumulation at the expense of beech seedlings, which showed significantly reduced growth in the mixture as compared to the monoculture. However, as this was only the case for Viburnum growing in the charcoal-filtered treatment, ozone seemed to counteract the beneficial effect of interspecific competition on above-ground biomass accumulation in Viburnum, while at the same time decreasing relative biomass allocation to roots. Foliar sensitivity of the two species was also altered under interspecific competition suggesting that results based on seedlings of single species grown in monocultures may significantly over- or under-estimate foliar sensitivity to ozone. These results demonstrate that competition is an important factor affecting plant responses to ozone stress, but the direction and severity of these effects depend on the interacting species.     

Do chronic aboveground O3 exposure and belowground pathogen stress affect growth and belowground biomass partitioning of juvenile beech trees (Fagus sylvatica L.)?

The impact of chronic free air ozone (O₃) exposure and belowground pathogen stress on growth and total biomass development of young beech trees (Fagus sylvatica L.) was investigated in a lysimeter study. Plants were growing during four years under ambient or elevated atmospheric O₃ concentrations. Additionally, in the last vegetation period the root rot pathogen Phytophthora citricola was introduced to study the interaction of ozone exposure and pathogen stress in the soil-plant system. A complete harvest at the end of the experiment enabled for the first time the assessment of fine and coarse root biomass of individual trees with a high vertical resolution down to two meter depth. Plant growth was significantly reduced by elevated ozone but not affected by P. citricola. Biomass partitioning between fine and coarse roots as well as vertical root distribution were significantly affected by both factors, whereas changes in root/shoot biomass ratio were not observed.     

UV-B辐射对窄叶野豌豆生长繁殖的影响

采用增补和滤除掉部分自然UV-B辐射的模拟试验,研究了增强和近环境UV-B辐射对高寒草甸一年生牧草窄叶野豌豆生长和繁殖的影响.结果表明:增补UV-B辐射处理后,窄叶野豌豆的株高、生物量、分配向果实的生物量、总花数和种子百粒重均显著下降,花期延迟,开花集中度和繁殖成功率有所提高,而种子产量无显著变化.相对于减弱UV-B辐射处理,近环境UV-B辐射使窄叶野豌豆的株高先降后升,分配向果实的生物量减少,花期、花数和种子产量无显著变化,种子百粒重减小.增强和近环境UV-B辐射对窄叶野豌豆的生长和繁殖有一定的抑制作用,且增强UV-B辐射的影响更大.     

Effects of species and soil‐nitrogen availability on root system architecture traits – study on a set of weed and crop species

Better managing crop : weed competition in cropping systems while reducing both nitrogen and herbicide inputs is a real challenge that requires a better understanding of crop and weed root architecture in relation to soil‐nitrogen availability. An original approach was used which considered the parameters of a simulation model of root architecture as traits to analyse (a) the interspecific diversity of root system architecture, and (b) its response to soil‐nitrogen availability. Two greenhouse experiments were conducted using three crop and nine weed species grown at two contrasted concentrations of soil‐nitrogen availability. Plant traits were measured to characterise both overall plant growth and root architecture, with a focus on primary root emergence, root elongation and branching. The studied root traits varied among species (from a twofold to a fourfold factor, depending on the trait), validating their use as indicators to analyse the interspecific variability of root architecture. The largest interspecies differences were for two traits: ‘maximal apical root diameter’ and ‘interbranch distance’ (distance between two successive laterals on the same root). Conversely, most of the studied root traits varied little with soil‐nitrogen availability (from no variation to a 1.1‐fold factor, depending on the trait) even though soil‐nitrogen availability varied with a 17‐fold factor and impacted the overall shoot and root biomass. So, the root traits used in this article are stable whatever soil‐nitrogen availability. As they reflect processes underlying root system architecture, this low effect of nitrogen suggests that the rules governing root architecture are little affected by plant nitrogen status and soil‐nitrogen availability. We propose that the determinants of differences in root system architecture between soils with contrasted nitrogen availability mainly originate from differences in the amount of carbon allocated to and within the root system. Characterising each plant species by a combination of root traits gave insights regarding the potential species competitive ability for soil resources in agroecosystems.     

Experimental setup of field lysimeters for studying effects of elevated ozone and below-ground pathogen infection on a plant-soil-system of juvenile beech (Fagus sylvatica L.)

An experiment, focusing on the effects of chronically enhanced O₃ regimes on young beech (Fagus sylvatica) and on the microbial rhizosphere community structure, was conducted from November 2002 to August 2006 in eight field lysimeters at the Helmholtz Zentrum München. The instrumentations of the lysimeters enabled the establishment of the water balance in the unsaturated zone and the assessment of the water uptake by plants. Further, the containment provided by the lysimeters made it possible to apply a root rot pathogen infection without contaminating the surrounding soil. A free-air fumigation system allowed to double the O₃ concentration in the air above four lysimeters relative to the ambient air. To avoid damage of the leaves the maximum O₃ concentration was limited to 150 nL L^?1. For nearly 70% of the time the set-point concentration was reached within 10%. In the final harvest the whole soil column was retrieved and a nearly complete data-set of above-ground and below-ground parameters became available.     

Contrasting ozone × pathogen interaction as mediated through competition between juvenile European beech (Fagus sylvatica) and Norway spruce (Picea abies)

Based on the growth-differentiation balance theory (GDB) and the influence of tropospheric ozone (O₃) on plants, we hypothesized that pre-conditioning with elevated O₃ reduces adverse effects of the root rot pathogen Phytophthora citricola Sawada. To this end a 2-year phytotron study with juvenile European beech (Fagus sylvatica L.) and (Picea abies [L.] Karst.) grown in mixture was performed. The hypothesis was tested on phenological, leaf and root morphological as well as physiological aspects of plant performance. Contrasting with spruce, elevated O₃ limited leaf and root biomass development, photosynthetic performance and N uptake of beech. The growth limitation by O₃ conveyed increased resistance in beech against the pathogen. Conversely, spruce displayed enhanced susceptibility in the combined O₃/P. citricola treatment. The hypothesis was supported in the case of beech rather than spruce. Nevertheless, conclusions support GDB regarding the trade-off between growth and stress defense, although compliance appears to be species-specific.     

Balancing positive and negative plant interactions: how mosses structure vascular plant communities

Our understanding of positive and negative plant interactions is primarily based on vascular plants, as is the prediction that facilitative effects dominate in harsh environments. It remains unclear whether this understanding is also applicable to moss–vascular plant interactions, which are likely to be influential in low-temperature environments with extensive moss ground cover such as boreal forest and arctic tundra. In a field experiment in high-arctic tundra, we investigated positive and negative impacts of the moss layer on vascular plants. Ramets of the shrub Salix polaris, herb Bistorta vivipara, grass Alopecurus borealis and rush Luzula confusa were transplanted into plots manipulated to contain bare soil, shallow moss (3 cm) and deep moss (6 cm) and harvested after three growing seasons. The moss layer had both positive and negative impacts upon vascular plant growth, the relative extent of which varied among vascular plant species. Deep moss cover reduced soil temperature and nitrogen availability, and this was reflected in reduced graminoid productivity. Shrub and herb biomass were greatest in shallow moss, where soil moisture also appeared to be highest. The relative importance of the mechanisms by which moss may influence vascular plants, through effects on soil temperature, moisture and nitrogen availability, was investigated in a phytotron growth experiment. Soil temperature, and not nutrient availability, determined Alopecurus growth, whereas Salix only responded to increased temperature if soil nitrogen was also increased. We propose a conceptual model showing the relative importance of positive and negative influences of the moss mat on vascular plants along a gradient of moss depth and illustrate species-specific outcomes. Our findings suggest that, through their strong influence on the soil environment, mat-forming mosses structure the composition of vascular plant communities. Thus, for plant interaction theory to be widely applicable to extreme environments such as the Arctic, growth forms other than vascular plants should be considered.