The effect of resource integration on fitness related traits in a clonal dune perennial,Hydrocotyle bonariensis

Summary The costs and benefits of resource integration in Hydrocotyle bonariensis were examined by comparing severed and intact clones grown across multiple resource gradients. Basipetal movement of water, nitrogen and photosynthates was demonstrated to occur between two rhizome branch systems interconnecting hundreds of ramets within a clone. Hydrocotyle clones of this size and larger have been shown to span highly patchy and unpredictable resource conditions in coastal dune environments. The extensive movement of water and nitrogen to portions of a clone deficient in these resources, resulted in a significant net benefit to the clone in terms of fitness-related traits: total biomass, ramet proliferation and seed production. The translocation of photosynthates across light gradients allowed for sexual reproduction and clonal expansion in the shade although this incurred a net cost to the clone in terms of ramet and seed production.     

CLONAL INTEGRATION ACROSS A SALT GRADIENT BY A NONHALOPHYTE,HYDROCOTYLE BONARIENSIS (APIACEAE)

This study examined the benefits associated with resource sharing among interconnected ramets spanning a soil salinity gradient. Clones of Hydrocotyle bonariensis, a rhizomatous dune perennial, expand into salt marsh communities from surrounding upland dune systems in coastal North Carolina. In rhizome-severing experiments conducted under both field and laboratory conditions, Hydrocotyle was shown to proliferate ramets under saline conditions, provided that these ramets were connected to other ramets growing in nonsaline conditions. Ramets that benefited from resource integration did not appear to be affected by local salt exposure in that these ramets were morphologically similar to those grown under nonsaline conditions. Supporting ramets incurred no net cost in terms of biomass or ramet production, but there was an increased percent allocation to roots and rhizomes. Ramets grown in saline conditions without the benefit of clonal integration showed high mortality and produced little or no net clonal growth. It is likely that the acropetal movement of water allowed Hydrocotyle clones to ameliorate the heterogeneous saline conditions associated with coastal environments.     

The effect of local resource availability and clonal integration on ramet functional morphology in Hydrocotyle bonariensis

Summary Within a physiologically integrated clone, the structure and functioning of an individual ramet is determined by: 1) the response of that ramet to its local environment and 2) its response to resource integration within the clone. In a multifactorial experiment, Hydrocotyle bonariensis ramets were grown in limiting resource environments with and without the benefit of basipetal resource movement from another branch of the clone. Ramets were analyzed for their morphological responses to variation in local light, water and nitrogen availability and to the superimposed effect of resource integration on these conditions. The expression of ramet morphology, from induction to development, was highly plastic in response to variable local resource availability. Resource integration changed a ramet's local response in a variety of ways depending on the resource(s) being translocated and the character involved. Among leaf characteristics (leaf weight, petiole height, blade area), resource translocation into the shade resulted in an enhancement of the local response. Similarly, the translocation of nitrogen and water generally increased clonal proliferation and sexual reproduction among ramets. In contrast, the translocation of water reversed the effect of local low water conditions on ramets by inhibiting root production. Some characters such as internode distance and leaf allometry were unaffected by integration. The maintenance of connections between ramets as a Hydrocotyle clone expands allows for resource sharing among widely separated ramets and can result in an integrated morpological response to a resource environment that is patchy in time and space.     

荒漠土壤因子和DSE定殖对克隆植物入侵的响应

克隆植物,尤其是游击型克隆植物,具有很强的扩展能力,通过克隆扩展可侵入到不同生境斑块。克隆植物入侵可能会影响入侵地土壤营养状况和微生物群落。为了探明克隆植物入侵对DSE(dark septate endophytes)活动和土壤理化性质的影响,于2013年6月在克隆植物羊柴(Hedysarum laeve)和沙鞭(Psammochloa villosa)群落空地沿根状茎延伸方向设置样方,分别于6月、8月和10月在样方内分0—10、10—20、20—30、30—40、40—50 cm土层采集土样和根样,研究了不同采样时间羊柴和沙鞭群落空地DSE和土壤理化性质时空变化。结果表明,从6月到10月,随时间后延,克隆植物逐渐侵入群落空地,沙鞭入侵群落空地数和分株数高于羊柴。羊柴群落空地根系DSE定殖率随采样时间后延,逐渐降低,最大值在6月;沙鞭群落空地根系DSE定殖率随采样时间后延,逐渐升高,最大值在10月。随着克隆植物入侵,入侵地土壤中可利用的营养物质含量显著提高,羊柴入侵提高了入侵群落空地土壤碱解N、有效P和速效K含量,沙鞭入侵提高了入侵群落空地土壤碱解N和有效P含量。相关性分析表明,羊柴群落空地DSE定殖率与土壤p H值和电导率显著正相关,沙鞭群落空地DSE定殖率与土壤p H值极显著负相关,与电导率、碱解N和有效P极显著正相关。克隆植物入侵使得土壤环境更有利于克隆植物自身生长,为荒漠植被恢复提供了前提。     

Spatial and temporal variation in islands of fertility in the Sonoran Desert

In many arid and semi-arid ecosystems, canopy trees and shrubs have a strong positive influence on soil moisture and nutrient availability, creating islands of fertility where organic matter and nutrients are high relative to areas outside the canopy. Previous studies of canopy effects on soil processes have rarely considered how landscape context may modulate these effects. We measured the effects of velvet mesquite trees (Prosopis velutina) on soil moisture and the biogeochemistry of nitrogen at different positions along a topographic gradient from upland desert to riparian zone in the Sonoran Desert of central Arizona. We also examined how landscape position and patterns of precipitation interact to determine the influence of P. velutina on soil moisture, N availability assessed using ion exchange resins, net N mineralization and net nitrification, and microbial biomass C and N. P. velutina clearly created islands of fertility with higher soil organic matter, net N mineralization and net nitrification rates, and microbial biomass under mesquite canopies. These effects were consistent across the landscape and showed little temporal variability. Magnitude and direction of effect of mesquite on soil moisture changed with landscape position, from positive in the upland to negative in the terrace, but only when soil moisture was >4%. Resin N showed responses to mesquite that depended on precipitation and topographic position, with highest values during wet seasons and under mesquite on terraces. We suggest changes in proximity of P. velutina to groundwater lead to shifts in biogeochemical processes and species interactions with change in landscape position along a topographic gradient.     

Laboratory experiments imply the conditionality of mycorrhizal benefits for Salix repens: role of pH and nitrogen to phosphorus ratios

Responses of one arbuscular mycorrhizal fungus (Glomus mosseae) and two ectomycorrhizal fungi (Hebeloma leucosarx, Paxillus involutus) to a range of substrate conditions were investigated in the laboratory. Non-mycorrhizal controls were also included. Substrate conditions included three levels of nitrogen – phosphorus ratios, ranging from N limitation to P limitation (N/P ratio 5.4, 16.2, 48.6), and three pH's, ranging from acidic to alkaline (pH 4, 5.5, 7), in a full factorial experiment. Plant parameters (carbon gain, N and P-content, root length) were significantly affected by fungus, soil pH and soil N/P, and their interactions. Mycorrhizal benefits by ectomycorrhizal fungi (EcMF) were generally larger than by the arbuscular mycorrhizal fungi (AMF), when assessed by above-ground parameters. Glomus mosseae, despite low colonization, had a much larger positive effect on root length than those EcMF. Hebeloma leucosarx and P. involutus were equally effective, despite differences in proportional colonization. Hebeloma leucosarx was able to expand niche width of S. repens towards alkaline conditions. Results are discussed in the framework of a dune successional gradient from young, calcareous, humus-poor towards old, acidic, humus-rich soils.     

Clonal integration facilitates the proliferation of smooth brome clones invading northern fescue prairies

Predicting exotic invaders and reducing their impacts on the biodiversity and function of native ecosystems require understanding of the mechanisms that facilitate their success during key stages of invasion. We determined whether clonal growth, characteristic of the majority of successful invaders of natural areas, facilitates the proliferation of Bromus inermis (smooth brome), an exotic grass invading prairie ecosystems across the Great Plains. By manipulating the below-ground connections of proliferating rhizomes as well as the levels of soil nitrogen along the margins of clones invading northern fescue prairies in Manitoba, Canada, we hypothesized that physiological integration would most benefit ramets invading low resource environments. Severing clonal connections reduced the mass of smooth brome shoots invading native prairies and was exacerbated by the immobilization of soil nutrients with glucose. Clonal connections were equally important in the maintenance of smooth brome density and the horizontal proliferation of ramets. Our results demonstrate the role of physiological integration in the proliferation of a clonal exotic invader and may help explain the success of clonal invaders in other regions. Although integration among invading ramets suggests several possibilities for successful management, future research must continue to elucidate differences in the invasiveness of native versus exotic species as well as the persistence of clonal connections among exotic invaders.     

Woody Plant Encroachment by <Emphasis Type="Italic">Juniperus virginiana</Emphasis> in a Mesic Native Grassland Promotes Rapid Carbon and Nitrogen Accrual

The cover and abundance of Juniperus virginiana L. in the U.S. Central Plains are rapidly increasing, largely as a result of changing land-use practices that alter fire regimes in native grassland communities. Little is known about how conversion of native grasslands to Juniperus-dominated forests alters soil nutrient availability and ecosystem storage of carbon (C) and nitrogen (N), although such land-cover changes have important implications for local ecosystem dynamics, as well as regional C and N budgets. Four replicate native grasslands and adjacent areas of recent J. virginiana encroachment were selected to assess potential changes in soil N availability, leaf-level photosynthesis, and major ecosystem C and N pools. Net N mineralization rates were assessed in situ over two years, and changes in labile soil organic pools (potential C and N mineralization rates and microbial biomass C and N) were determined. Photosynthetic nitrogen use efficiencies (PNUE) were used to examine differences in instantaneous leaf-level N use in C uptake. Comparisons of ecosystem C and N stocks revealed significant C and N accrual in both plant biomass and soils in these newly established forests, without changes in labile soil N pools. There were few differences in monthly in situ net N mineralization rates, although cumulative annual net N mineralization was greater in forest soils compared to grasslands. Conversely, potential C mineralization was significantly reduced in forest soils. Encroachment by J. virginiana into grasslands results in rapid accretion of ecosystem C and N in plant and soil pools with little apparent change in N availability. Widespread increases in the cover of woody plants, like J. virginiana, in areas formerly dominated by graminoid species suggest an increasing role of expanding woodlands and forests as regional C sinks in the central U.S.     

Environmental heterogeneity and clonal growth: a study of the capacity for reciprocal translocation in Glechoma hederacea L.

Clonal fragments of Glechoma hederacea L. (Lamiaceae) were subjected to environments in which light and nutrients were supplied with a strictly negative association in space, i.e. when one of these resources was in ample supply the other was scarce. Treatments were chosen to simulate environments in which clones grew either within homogeneous conditions or across patch types (heterogeneous conditions). The hypothesis was tested that reciprocal translocation (i.e. exchange of both nutrients and assimilates) between connected groups of ramets would increase biomass production of clones growing under heterogeneous conditions compared to that of clones growing in homogeneous conditions. A cost-benefit analysis was carried out to test this hypothesis. Results suggested that reciprocal translocation did not occur at the structural scale considered in this experiment; no evidence was found for a significant effect on whole clone biomass of assimilate and/or nutrient translocation between clone parts experiencing contrasting levels of resource supply. It is suggested that predominantly acropetal movement of resources and the pattern of integrated physiological unit formation in G. hederacea are the main properties responsible for the lack of mutual physiological support between connected clonal fragments growing in differing habitat conditions. These properties are expected to promote clonal expansion and the exploitation of new territory, rather than sustaining clone parts in sub-optimal patches of habitat for prolonged periods of time.     

Foliar Nitrogen Dynamics and Nitrogen Resorption of a Sandy Shrub Salix gordejevii in Northern China

Resorption of nitrogen (N) from senescing leaves is an important conservation mechanism that allows plants to use the same N repeatedly. Seasonal variations in leaf nitrogen of mature green and senescing leaves and N resorption in Salix gordejevii Chang, a sandy shrub in northern China, were studied. Our objective was to compare N resorption of this Salix species that successfully occupy different habitats (shifting sandland, fixed sandland and lowland) with differences in soil N availability and moisture. Nitrogen concentrations in green and senescing leaves were higher in June and July. N resorption efficiency (percentage reduction of N between green and senescing leaves) was highest at shifting sandland, intermediate at fixed sandland, and lowest at lowland. There was a clear seasonal variation in N-resorption efficiency, with a lower value at the early growing season and a higher value during summer. N resorption efficiency was lower at the sites with higher soil N availability, suggesting that the efficiency of the resorption process is determined by the availability of the nutrient in the soil. Resorption from senescing leaves may play an important role in the nitrogen dynamics of sandy plants and reduce the nitrogen requirements for plant growth. We conclude that N resorption from senescing leaves in S. gordejevii was correlated to soil characteristics and higher N resorption on poor soils is a phenotypic adjustment by this species to maximize N-use at low availability.     

Nitrogen translocation between clonal mother and daughter trees at a grassland–forest boundary

There is abundant evidence from short-term experiments using herbs that nutrients can be translocated from mother ramets to daughter ramets, but there is little long-term evidence from woody plants. Here, we examine translocation in field populations of a clonal tree over two growing seasons. We applied ¹⁵N to mothers or daughters in clones of Populus tremuloides at the northern edge of the North American Great Plains, where mother ramets form closed-canopy stands on relatively nutrient-rich soils, and daughter ramets occur nearby in relatively nutrient-poor grasslands. Unlabeled daughters in clones with labeled mothers had δ¹⁵N values significantly greater than those in unlabeled clones, confirming translocation from mothers to daughters. However, unlabeled mothers in clones with labeled daughters also had δ¹⁵N values significantly greater than those in unlabeled clones, indicating translocation from daughters to mothers. Further, the total foliage accumulation of added ¹⁵N was significantly (c. 10×) greater in mothers than in daughters, suggesting that more N was translocated from daughters to mothers, than from mothers to daughters. Thus, ¹⁵N moved both from mothers to daughters and from daughters to mothers, with net flow toward mothers. Because long-lived woody ramets in the field face nutrient competition from other ramets, interspecific neighbors, and soil microbes, the environmental availability of nutrients for uptake may be low for both mother and daughter ramets, causing translocation within a clone to be toward larger ramets with greater demand.     

Clonal integration is beneficial for resource sharing in a creeping amphibian herb (<Emphasis Type="Italic">Alteranthera philoxeroides</Emphasis>)

Clonal integration facilitates the growth and reproduction of clonal plants by providing the ability to share resources among ramets in heterogeneous environments. The benefits of clonal integration for plant growth may depend on a contrast in resource availability and may encounter costs, especially when a young part of the clone is growing across a border between richer and poorer conditions than the old part. We studied a clonal amphibian plant growing across a border between an aquatic and a terrestrial ecosystem, which typically differ in the availability of resources. We asked whether the young part of the clone is supporting the old part with phosphorus and whether this support has costs. We performed an experiment with Alternanthera philoxeroides where plants grow from water to a terrestrial habitat. The terrestrial habitat had either a low or high phosphorus supply, and the connection between the old and young parts of the clone was either left intact or split. We determined that the young part of the clone growing in a terrestrial habitat supported the old part with phosphorus when growing on a substrate rich in phosphorus. We have found no cost of this resource translocation; on the contrary, whole clones increased not only their accumulation of phosphorus, but also of nitrogen. Our study shows how an amphibian plant may profit from heterogeneous habitats by resource sharing in a clonal network.     

Root decomposition across a barrier island chronosequence: Litter quality and environmental controls

A root decomposition study using the litterbag approach was conducted along a dune and swale chronosequence on the Virginia Coast Reserve-Long Term Ecological Research Site in Virginia, USA to evaluate how environmental and substrate quality factors influence belowground decay and associated nutrient dynamics. Gradients in moisture levels and nitrogen availability associated with the chronosequence provided the experimental framework. Spartina patens roots were buried at all sites as a standard substrate to evaluate environmental influences. Roots native to each site were buried to evaluate community decay dynamics and the influence of litter quality. Spartina decay was reduced in the wet, anoxic soils of swale sites (k = 0.21–0.33 yr^-1) relative to decay in dunes soils (k = 0.52–0.72 yr^-1). Increasing soil nitrogen availability from younger to older sites had no effect on the rate of Spartina root decay. Native root decay across the Hog Island chronosequence exhibits certain trends expected in response to nitrogen limitation and moisture availability. Increased nitrogen content of root material corresponds to increased soil nitrogen availability. Among dune sites, native root decay increased in concert with increased root nitrogen (6 year k = 0.34 yr^-1, 120 year dune: k = 0.97 yr^-1). Litter quality, alone, does not explain this trend since Spartina roots decayed more slowly than native dune roots and had a higher initial nitrogen content. Among swales, increased moisture levels and associated soil anoxia inhibited native root decomposition and minimized the effects of litter quality on decay. In general, phosphorus was rapidly lost from decaying roots while nitrogen immobilization was low to nonexistent. The low nitrogen immobilization of decaying roots in a nitrogen limited ecosystem warrants further study and may reveal that belowground decay increases the rate of nutrient cycling relative to decay aboveground.     

Effect of mineral nitrogen on nitrogen nutrition and biomass partitioning between the shoot and roots of pea (Pisum sativum L.).

The effect of mineral N availability on nitrogen nutrition and biomass partitioning between shoot and roots of pea (Pisum sativum L., cv Baccara) was investigated under adequately watered conditions in the field, using five levels of fertiliser N application at sowing (0, 50, 100, 200 and 400 kg N ha^–1). Although the presence of mineral N in the soil stimulated vegetative growth, resulting in a higher biomass accumulation in shoots in the fertilised treatments, neither seed yield nor seed nitrogen concentration was affected by soil mineral N availability. Symbiotic nitrogen fixation was inhibited by mineral N in the soil but it was replaced by root mineral N absorption, which resulted in optimum nitrogen nutrition for all treatments. However, the excessive nitrogen and biomass accumulation in the shoot of the 400 kg N ha^–1 treatment caused crop lodging and slightly depressed seed yield and seed nitrogen content. Thus, the presumed higher carbon costs of symbiotic nitrogen fixation, as compared to root mineral N absorption, affected neither seed yield nor the nitrogen nutrition level. However, biomass partitioning within the nodulated roots was changed. The more symbiotic nitrogen fixation was inhibited, the more root growth was enhanced. Root biomass was greater when soil mineral N availability was increased: root growth was greater and began earlier for plants that received mineral N at sowing. Rooting density was also promoted by increased mineral N availability, leading to more numerous but finer roots for the fertilised treatments. However, the maximum rooting depth and the distribution of roots with depth were unchanged. This suggested an additional direct promoting effect of mineral N on root proliferation.     

Effects of inorganic nitrogen and phosphorus supply on nitrogen aquisition from alanine by birch seedlings in symbiosis with Paxillus involutus

Northern forests are exposed to relatively high ammonia inputs due to high atmospheric deposition and the common practise of forest fertilization. It is not known how increased soil NH₄ ⁺concentrations affect acquisition of symbiosis-mediated N from organic sources. We examined the effect of inorganic N and P availability on N acquisition from alanine by 43 weeks old birch (Betula pendula) seedlings in symbiosis with the ectomycorrhizal fungus Paxillus involutus. The seedlings were exposed for 9 weeks to nutrient additions equivalent to 43 kg N and 6.4 kg P ha^-1 (low N and P availability), 250 kg N and 38 kg P ha^-1(high N and P availability) or to 250 kg N and 6.4 kg P ha^-1 (high N and low P availability). Carbon and nitrogen allocation between the symbionts was assessed by exposing the foliage to ¹⁴CO₂ and the mycelium to ¹⁵N-alanine or ¹⁵NH₄ ⁺ simultaneously and measuring the distribution of the isotopic tracers after a three-day chase period. High inorganic N combined with low P availability did not have marked effect on symbiosis-mediated N uptake from alanine, whilst high N and P availability reduced alanine-derived ¹⁵N translocation by the fungus to the plant. Shoot ¹⁵N concentration and concentration of ¹⁴C in the extramatrical mycelium correlated significantly across treatments pointing to controlled reciprocity of transactions between the partners.     

Root nodulation and nitrogen accumulation and partitioning in legume crops as affected by soil solarization

Soil solarization is a preplanting technique used in hot climates to control weeds and soilborne pathogens consisting of mulching the soil surface with polyethylene sheets. The increase in temperature associated with solarized soil could affect nitrogen availability for grain legume crops through effects on nitrogen fixing soil microorganisms or other mechanisms. To examine the effects of solarization on natural root nodulation and nitrogen accumulation and partitioning in the plant, two solarization field experiments were carried out over two planting seasons, involving genotypes of both faba bean (Vicia faba) and chickpea (Cicer arietinum). The effect of sowing date was also studied in the first season. Solarization increased the maximum soil temperature by 9–10 °C in the first, and by 13–15 °C in the second season. At 5 cm below the solarized soil surface, a temperature of over 46 °C prevailed for 146 and 280 h over the two respective seasons, while this temperature was not attained in unmulched soil. Solarization delayed the initiation of nodulation and consistently reduced the nodule number per host plant, but generated an approximate doubling of mean nodule weight. The total nodule mass per plant was not affected by the treatment in the first season, but was reduced in the second season. Solarization significantly increased the concentrations of NO₃^– -N, Na⁺, Zn²⁺, Ca²⁺ and K⁺ in the soil extract, and the total nitrogen accumulated in the whole plant. This latter increase was due to both higher plant growth and a greater plant nitrogen concentration. The increased nitrogen level in the plant was not uniform with respect to plant component, varying from 57% in the roots to 198% in the pods and seeds. The plants grown in non-solarized soil accumulated about 31% of their total N content in the shoots of the parasitic weed Orobanche crenata. Solarization dramatically improved grain yield by 300–900% in both seasons and in all genotypes studied, due to increased N availability in soil, N accumulation in plants, improved plant growth, and complete control of the parasite weed O. crenata. On the basis of these beneficial effects, soil solarization, which avoids site contamination and is suited to organic farming, should be a good opportunity in Mediterranean areas where the level and stability of grain yields are low, and the infestation of O. crenata is high.     

Clonal integration in Fragaria chiloensis differs between populations: ramets from grassland are selfish

In plants, only species with clonal growth are able to directly transfer resources between otherwise independent units of the same genetic individual. A simple conceptual model of plant performance as a function of internal resource supply and environmental resource availability suggests that resource sharing between ramets within clones is likely to be disadvantageous in uniform habitats and advantageous in patchy ones. It was therefore hypothesized that clones in populations from relatively uniform habitats will have been selected for low rates of resource sharing between ramets compared to clones in populations from patchier habitats. In coastal northern California, the clonal herb Fragaria chiloensis is common both in grasslands, where resources are relatively uniform, and on sand dunes, where resources are more patchy. It was predicted that clones from a grassland population of Fragaria would have “selfish” ramets with low rates of resource sharing compared to clones from an adjacent dune population. Ramets were subjected to contrasting light levels with and without connection between ramets. Patterns of biomass accumulation were consistent with the prediction. This appears to be the first report of genetically based variation in patterns of resource sharing in clonal plants. It supports the idea that these patterns are locally selected to increase plant performance in habitats with different patterns of resource availability. Received: 19 August 1998 / Accepted: 4 March 1999     

The effects of clonal integration on morphological plasticity and placement of daughter ramets in black locust (Robinia pseudoacacia)

We studied the field response of Robinia pseudoacacia L. to light, total soil nitrogen, available soil phosphorus and soil pH. Results indicated that there was very strong clonal integration between mother and daughter ramets. Mother ramets can provide nitrogen and phosphorus to daughter ramets sufficient for their continued growth through strong clonal integration, but cannot provide enough photosynthate. With clonal integration, soil nitrogen and phosphorus availability had no effect on biomass allocation to roots, number of ramets and length of connection roots. Biomass allocation to roots increased markedly and responded to nitrogen and phosphorus availability, when the connections were severed. Light had a significant effect on the percent of biomass allocation to leaves and number of ramets, but no effect on the length of connection roots. Daughter ramets allocated more resources to leaves, and clones placed more daughter ramets in high light patches than in low light patches. Soil pH had a significant effect on ramet number and connection root length. Clones concentrated in alkaline patches and escaped from acid patches through selective placement of daughter ramets and changing the length of connection roots. We suggest that the clonal integration may be very strong and provide sufficient soil resources to daughter ramets, then affect the daughter ramets’ morphology and placement, if the size of a specific ramet is significantly larger than the other ramets in an arbor clone.     

Plasticity in R/S ratio, morphology and fitness-related traits in response to reciprocal patchiness of light and nutrients in the stoloniferous herb, Glechoma longituba L

Clonal fragments of the stoloniferous herb Glechoma longituba were subjected to a complementary patchiness of light and soil nutrients including two spatially homogeneous treatments (SR–SR and IP–IP) and two spatially heterogeneous treatments (IP–SR and SR–IP). SR and IP indicate patches (shaded, rich) with low light intensity (shaded, S), high nutrient availability (rich, R) and patches (illuminated, poor) with high light intensity (illuminated, I) and low nutrient availability (poor, P), respectively. Plasticity of the species in root–shoot ratio, fitness-related traits (biomass, number of ramets and dry weight per ramet) and clonal morphological traits (length and specific length of stolon internodes, area and specific area of laminae, length and specific length of petioles) were experimentally examined. The aim is to understand adaptation of G. longituba to the environment with reciprocal patches of light and soil nutrients by plasticities both in root–shoot ratio and in (clonal) morphology. Our experiment revealed performance of the clonal fragments growing from patches with high light intensity and low soil nutrient availability into the adjacent opposite patches was increased in terms of the fitness-related characters. R/S ratio and clonal morphology were plastic. Meanwhile, the capture of light resource from the light-rich patches was enhanced while the capture of soil nutrients from either the nutrient-rich or the nutrient-poor patches was not. Analysis of cost and benefit disclosed positive effects of clonal integration on biomass production of ramets in the patches with low light intensity and high soil nutrient availability. These results suggest an existence of reciprocal translocation of assimilates and nutrients between the interconnected ramets. The reinforced performance of the clonal fragments seems to be related with specialization of clonal morphology in the species.     

Growth and allocation of the arctic sedges Eriohorum angustifolium and E. vaginatum: effects of variable soil oxygen and nutrient availability

In arctic tundra soil, oxygen depletion associated with soil flooding may control plant growth either directly through anoxia or indirectly through effects on nutrient availability. This study was designed to evaluate whether plant growth and physiology of two arctic sedge species are more strongly controlled by the direct or indirect effects of decreased soil aeration. Eriophorum angustifolium and E. vaginatum, which originate from flooded and well-drained habitats, respectively, were grown in an in situ transplant garden at two levels of soil oxygen, nitrogen, and phosphorus availability over two growing seasons. In both species, N addition had a stronger effect on growth and biomass allocation than P addition or soil oxygen depletion. Net photosynthesis and carbohydrate concentrations were relatively insensitive to changes in these factors. Biomass reallocated from shoots to below-ground parts in response to limited N supply was equally divided between roots (nutrient acquisition) and perennating rhizomes (storage tissue formation) in E. angustifolium. E. Vaginatum only increased its allocation to rhizomes. In the flood-tolerant E. angustifolium, growth was improved by soil anoxia and biomass allocation among plant parts was not significantly affected. Contrary to our initial hypothesis, whole-plant growth in E. vaginatum improved in flooded soils; however, it only did so when N availability was high. Under low N availability growth in flooded soils was reduced by 20% compared to growth in the aerobic environment. Reduced biomass allocation to rhizomes and thus to storage potential under anaerobic conditions may reduce long-term survival of E. vaginatum in flooded habitats.