Biotic contexts alter metal sequestration and AMF effects on plant growth in soils polluted with heavy metals

Investigating how arbuscular mycorrhizal fungi (AMF)-plant interactions vary with edaphic conditions provides an opportunity to test the context-dependency of interspecific interactions. The relationship between AMF and their host plants in the context of other soil microbes was studied along a gradient of heavy metal contamination originating at the site of zinc smelters that operated for a century. The site is currently under restoration. Native C3 grasses have reestablished, and C4 grasses native to the region but not the site were introduced. Interactions involving the native mycorrhizal fungi, non-mycorrhizal soil microbes, soil, one C3 grass (Deschampsia flexuosa), and one C4 grass (Sorghastrum nutans) were investigated using soils from the two extremes of the contamination gradient in a full factorial greenhouse experiment. After 12 weeks, plant biomass and root colonization by AMF and non-mycorrhizal microbes were measured. Plants from both species grew much larger in soil from low-contaminated (LC) origin than high-contaminated (HC) origin. For S. nutans, the addition of a non-AMF soil microbial wash of either origin increased the efficacy of AMF from LC soils but decreased the efficacy of AMF from HC soils in promoting plant growth. Furthermore, there was high mortality of S. nutans in HC soil, where plants with AMF from HC died sooner. For D. flexuosa, plant biomass did not vary with AMF source or the microbial wash treatment or their interaction. While AMF origin did not affect root colonization of D. flexuosa by AMF, the presence and origin of AMF did affect the number of non-mycorrhizal (NMF) morphotypes and NMF root colonization. Adding non-AMF soil biota reduced Zn concentrations in shoots of D. flexuosa. Thus the non-AMF biotic context affected heavy metal sequestration and associated NMF in D. flexuosa, and it interacted with AMF to affect plant biomass in S. nutans. Our results should be useful for improving our basic ecological understanding of the context-dependency of plant-soil interactions and are potentially important in restoration of heavy-metal-contaminated sites.     

Effects of soil biota from different ranges on Robinia invasion: acquiring mutualists and escaping pathogens

The net effects of soil biota on exotic invaders can be variable, in part, because net effects are produced by many interacting mutualists and antagonists. Here we compared mutualistic and antagonistic biota in soils collected in the native, expanded, and invasive range of the black locust tree, Robinia pseudoacacia. Robinia formed nodules in all soils with a broad phylogenetic range of N-fixing bacteria, and leaf N did not differ among the different sources of soil. This suggests that the global expansion of Robinia was not limited by the lack of appropriate mutualistic N-fixers. Arbuscular mycorrhizal fungi (AMF) from the native range stimulated stronger positive feedbacks than AMF from the expanded or invasive ranges, a biogeographic difference not described previously for invasive plants. Pythium taxa collected from soil in the native range were not more pathogenic than those from other ranges; however, feedbacks produced by the total soil biota were more negative from soils from the native range than from the other ranges, overriding the effects of AMF. This suggests that escape from other pathogens in the soil or the net negative effects of the whole soil community may contribute to superior performance in invaded regions. Our results suggest that important regional evolutionary relationships may occur among plants and soil biota, and that net effects of soil biota may affect invasion, but in ways that are not easily explained by studying isolated components of the soil biota.     

Are savannas patch-dynamic systems? A landscape model

Savannas are ecosystems characterized by the coexistence of woody species (trees and bushes) and grasses. Given that savanna characteristics are mainly formed from competition, herbivory, fire, woodcutting, and patchy soil and precipitation characteristics, we propose a spatially explicit model to examine the effects of the above-mentioned parameters on savanna vegetation dynamics in space and time. Furthermore, we investigate the effects of the above-mentioned parameters on tree–bush–grass ratios, as well as the degrees of aggregation of tree–bush–grass biomass. We parameterized our model for an arid savanna with shallow soil depth as well as a mesic one with generally deeper and more variable soil depths. Our model was able to reproduce savanna vegetation characteristics for periods of time over 2000 years with daily updated time steps. According to our results, tree biomass was higher than bush biomass in the arid savanna but bush biomass exceeded tree and grass biomass in the simulated mesic savanna. Woody biomass increased in our simulations when the soil's porosity values were increased (mesic savanna), in combination with higher precipitation. Savanna vegetation varied from open savanna to woodland and back to open savanna again. Vegetation cycles varied over ∼300-year cycles in the arid and ∼220-year cycles in the mesic-simulated savanna. Autocorrelation values indicated that there are both temporal and spatial vegetation cycles. Our model indicated cycling savanna vegetation at the landscape scale, cycles in cells, and patchiness, i.e. patch dynamics.     

The invasive grass Agropyron cristatum doubles belowground productivity but not soil carbon

Root dynamics are among the largest knowledge gaps in determining how terrestrial carbon (C) cycles will respond to environmental change. Increases in productivity accompanying plant invasions and introductions could increase ecosystem C storage, but belowground changes are unknown, even though roots may account for 50-90% of production in temperate ecosystems. We examined whether the introduction of a widespread invasive grass with relatively high shoot production also increased belowground productivity and soil C storage, using a multiyear rhizotron study in 50-year-old stands dominated either by the invasive C3 grass Agropyron cristatum or by largely C4 native grasses. Relative to native vegetation, stands dominated by the invader had doubled root productivity. Soil carbon isotope values showed that the invader had made detectable contributions to soil C. Soil C content, however, was not significantly different between invader-dominated stands (0.42 mg C/g soil) and native vegetation (0.45 mg C/g soil). The discrepancy between enhanced production and lack of soil C changes was attributable to differences in root traits between invader-dominated stands and native vegetation. Relative to native vegetation, roots beneath the invader had 59% more young white tissue, with 80% higher mortality and 19% lower C:N ratios (all P < 0.05). Such patterns have previously been reported for aboveground tissues of invaders, and we show that they are also found belowground. If these root traits occur in other invasive species, then the global phenomenon of increased productivity following biological invasion may not increase soil C storage.     

Plant winners and losers during grassland N-eutrophication differ in biomass allocation and mycorrhizas

Human activities release tremendous amounts of nitrogenous compounds into the atmosphere. Wet and dry deposition distributes this airborne nitrogen (N) on otherwise pristine ecosystems. This eutrophication process significantly alters the species composition of native grasslands; generally a few nitrophilic plant species become dominant while many other species disappear. The functional equilibrium model predicts that, compared to species that decline in response to N enrichment, nitrophilic grass species should respond to N enrichment with greater biomass allocation aboveground and reduced allocation to roots and mycorrhizas. The mycorrhizal feedback hypothesis states that the composition of mycorrhizal fungal communities may influence the composition of plant communities, and it predicts that N enrichment may generate reciprocal shifts in the species composition of mycorrhizal fungi and plants. We tested these hypotheses with experiments that compared biomass allocation and mycorrhizal function of four grass ecotypes (three species), two that gained and two that lost biomass and cover in response to long-term N enrichment experiments at Cedar Creek and Konza Long-Term Ecological Research grasslands. Local grass ecotypes were grown in soil from their respective sites and inoculated with whole-soil inoculum collected from either fertilized (FERT) or unfertilized (UNFERT) plots. Our results strongly support the functional equilibrium model. In both grassland systems the nitrophilic grass species grew taller, allocated more biomass to shoots than to roots, and formed fewer mycorrhizas compared to the grass species that it replaced. Our results did not fully support the hypothesis that N-induced changes in the mycorrhizal fungal community were drivers of the plant community shifts that accompany N eutrophication. The FERT and UNFERT soil inoculum influenced the growth of the grasses differently, but this varied with site and grass ecotype in both expected and unexpected ways suggesting that ambient soil fertility or other factors may be interacting with mycorrhizal feedbacks.     

丛枝菌根对土壤-植物系统中重金属迁移转化的影响

丛枝菌根真菌(arbuscular mycorrhizal fungi, AMF)是一类在自然和农业生态系统中广泛存在并能与多数陆生植物形成共生关系的土壤真菌,在重金属污染土壤中对宿主植物的生长及吸收累积重金属具有重要影响,因而对污染土壤的生物修复具有潜在应用价值。以重金属从根际土壤进入植物并在植物体内再分配过程为主线,介绍丛枝菌根在这一过程中对重金属环境行为,特别是根际土壤中重金属赋存形态及植物吸收重金属的影响。最后,对丛枝菌根影响植物重金属耐性机制研究前沿和菌根修复技术的应用前景进行展望。     

喀斯特生境下AMF侵染对任豆生长的影响

喀斯特地区土壤退化,植被定植更新困难,丛枝菌根真菌(Arbuscular mycorrhizal fungi,AMF)具有增强植物养分吸收能力和抵抗逆境胁迫能力。研究喀斯特生境下植物与AMF共生效果,选择优势菌种促进喀斯特植被恢复,对于提高植物定植成活率具有重要作用。以豆科植物任豆(Zenia insignis)幼苗为试验材料,盆栽条件下,选取喀斯特优势菌种-摩西球囊霉(Funneliformis mosseae)、根内球囊霉(Rhizophagus intraradices),2种菌根真菌混合菌剂进行接种,培养180 d,研究贫瘠喀斯特土壤生境和养分较高的滇柏林下土壤生境下AMF对任豆生长影响。结果表明:摩西球囊霉、根内球囊霉和混合接种均能侵染任豆根系,幼嫩根系更易侵染,木质化根系侵染率下降。接种摩西球囊霉,贫瘠喀斯特土壤生境下,株高、地径、地上生物量、地下生物量和总生物量分别提高68.92%、56.18%、83.90%、42.20%和67.34%;养分较高的滇柏林下喀斯特土壤生境下,株高、地上生物量、地下生物量和总生物量分别提高48.05%、6.77%、7.92%和8.89%;根内球囊霉处理接种效应低于摩西球囊霉和混合接种处理,对生物量增长为负效应,混合接种处理接种效应介于单接种之间,摩西球囊霉接种效果优于根内球囊霉和混合接种。摩西球囊霉在贫瘠喀斯特土壤生境下发挥的促生效应优于养分较高的喀斯特土壤,可作为喀斯特侵蚀区植被恢复菌根真菌干扰途径的优势菌种,混合接种作为接种剂具有单接种兼容效应。     

Additive effects of aboveground polyphagous herbivores and soil feedback in native and range-expanding exotic plants

Plant biomass and plant abundance can be controlled by aboveground and belowground natural enemies. However, little is known about how the aboveground and belowground enemy effects may add up. We exposed 15 plant species to aboveground polyphagous insect herbivores and feedback effects from the soil community alone, as well as in combination. We envisaged three possibilities: additive, synergistic, or antagonistic effects of the aboveground and belowground enemies on plant biomass. In our analysis, we included native and phylogenetically related range-expanding exotic plant species, because exotic plants on average are less sensitive to aboveground herbivores and soil feedback than related natives. Thus, we examined if lower sensitivity of exotic plant species to enemies also alters aboveground-belowground interactions. In a greenhouse experiment, we exposed six exotic and nine native plant species to feedback from their own soil communities, aboveground herbivory by polyphagous insects, or a combination of soil feedback and aboveground insects and compared shoot and root biomass to control plants without aboveground and belowground enemies. We observed that for both native and range-expanding exotic plant species effects of insect herbivory aboveground and soil feedback added up linearly, instead of enforcing or counteracting each other. However, there was no correlation between the strength of aboveground herbivory and soil feedback. We conclude that effects of polyphagous aboveground herbivorous insects and soil feedback add up both in the case of native and related range-expanding exotic plant species, but that aboveground herbivory effects may not necessarily predict the strengths of soil feedback effects.     

Fungal endophytes directly increase the competitive effects of an invasive forb

Competitive outcomes among plants can vary in different abiotic and biotic conditions. Here we tested the effects of two phylotypes of Alternaria endophytes on the growth, competitive effects, and competitive responses of the exotic invasive forb Centaurea stoebe. Centaurea stoebe was a better competitor against North American grass species than grasses from its European home range in the absence of endophytes. However, one endophyte both increased the biomass of C. stoebe and reduced the competitive effect of North American grasses on C. stoebe. The competitive effects of C. stoebe on grass species native to North America were enhanced by both fungal endophytes, but not for native European grasses. We do not know the mechanism by which endophytes increased C. stoebe's competitive ability, and particularly against biogeographically new neighbors, but one endophyte increased the competitive ability of C. stoebe without increasing its size, suggesting mechanisms unrelated to increased growth. We tested only a fraction of the different endophytic fungi that have been found in C. stoebe, only scratching the surface of understanding their indirect effects. However, our results are the first to demonstrate such effects of a fungal endophyte infecting an invasive forb, and one of the few to show that endophyte effects on competition do not have to be mediated through herbivory.     

Are soil mite assemblages structured by the identity of native and invasive alien grasses?

Associations between plants and animals in aboveground communities are often predictable and specific. This has been exploited for the purposes of estimating the diversity of animal species based on the diversity of plant species. The introduction of invasive alien plants into an ecosystem can result in dramatic changes in both the native plant and animal assemblages. Few data exist at the species level to determine whether belowground animal assemblages share the same degree of association to plants. The hypotheses that soil mites (Acari) form assemblages specifically associated with different native grass species in an unmanipulated natural ecosystem and that invasive alien grasses will impact soil mite assemblage composition in this setting were tested. Soil mites sampled beneath five native and two invasive alien species of grasses at the Konza Prairie Biological Station, Kansas, USA, were similarly abundant, species rich, diverse, and taxonomically distinct. No mite species had affinities for a specific grass species. There was no evidence from analysis of similarity, canonical correspondence analysis, or a nonparametric assemblage analysis that the assemblage composition of soil mites was specific to grass species. Results suggest that soil mite assemblages were more related to characteristics of the plant assemblage as a whole or prevailing soil conditions. The most recent invasive alien grass did not support a successionally younger mite fauna, based on the ratio of mesostigmatid to oribatid mites, and neither of the two invasive grasses influenced mite assemblage structure, possibly because they had not yet substantially altered the soil environment. Our results suggest that extrapolations of soil mite diversity based on assumptions of plant specificity would be invalid.     

Role of arbuscular mycorrhizal fungi on the performance of floodplain Phragmites japonica under nutrient stress condition

A pot experiment was conducted to evaluate the potential effects of arbuscular mycorrhizal fungi (AMF) on growth, nutrient uptake, and inoculation effectiveness on Phragmites japonica. Spores of AMF strains (Gigaspora margarita Becker &; Hall) were collected from the commercial product ‘Serakinkon’. Four treatments, namely, natural soil (NS), natural soil inoculated by AM fungi, sterilised soil (SS) inoculated by AM fungi, and SS without AM fungi inoculation were selected to determine the effects of applied and indigenous AMF on P. japonica. The average colonisation level of P. japonica was 24–33%, whereas no colonisation was found in the SS. AMF colonisation increased the chlorophyll content (r?=?0.84, p?r?=?0.89, p?相似文献   

Plant species differ in their ability to reduce allocation to non-beneficial arbuscular mycorrhizal fungi

Theory suggests that cheaters threaten the persistence of mutualisms, but that sanctions to prevent cheating can stabilize mutualisms. In the arbuscular mycorrhizal symbiosis, reports of parasitism suggest that reductions in plant carbon allocation are not universally effective. I asked whether plant species differences in mycorrhizal responsiveness would affect both their susceptibility to parasitism and their reduction in allocation to non-beneficial arbuscular mycorrhizal fungi (AMF) in high-phosphorus soils. In a greenhouse experiment, I found that two C3 grasses, Bromus inermis and Elymus repens, effectively suppressed root colonization and AMF hyphal abundance. Increases in soil phosphorus did not reduce the degree to which AMF increased plant biomass. In contrast, two C4 grasses, Andropogon gerardii and Schizachyrium scoparium, more weakly reduced root colonization and failed to suppress AMF hyphal abundance. Consequently, they experienced strong declines in their response to AMF, and one species suffered parasitism. Thus, species differ in susceptibility to parasitism and their reduction in allocation to non-beneficial AMF. These differences may affect the distribution and abundance of plants and AMF, as well as the stability of the mutualism.     

Grass feedbacks on fire stabilize savannas

Savannas commonly consist of a discontinuous cover of overstory trees and a groundcover of grasses. Savanna models have previously demonstrated that vegetation feedbacks on fire frequency can limit the density of overstory trees, thereby maintaining savannas. Positive feedbacks of either savanna trees alone or trees and grasses together on fire frequency have been shown to result in a stable savanna equilibrium. Grass feedbacks on fire frequency, in contrast, have resulted in stable equilibria in either a grassland or forest state, but not in a savanna. These results, however, were derived from a system of differential equations that assumes that fire occurrence is strictly deterministic and that vegetation losses due to fire are continuous in time. We develop an alternative formulation of the grass-fire feedback model that assumes that fires are discrete and occur stochastically in time to examine the influence of these assumptions on the predicted state of the system. We show that incorporating fire as a discrete event can produce a recurring temporal refuge in which both grass and trees co-occur in a stable, bounded savanna. In our model, tree abundance is limited without invoking demographic bottlenecks in the transition from fire-sensitive to fire-resistant life history stages. An increasing strength of grass feedback on fire results in regular, predictable fires, which suggests that the system can also be modeled using a set of difference equations. We implement this discrete system using modified Leslie/Gower difference equations and demonstrate the existence of a bounded savanna state in this model framework. Our results confirm the potential for grass feedbacks to result in stable savannas, and indicate the importance of modeling fire as a discrete event rather than as a loss rate that is continuous in time.     

Mycorrhizal fungal identity and diversity relaxes plant-plant competition

There is a great interest in ecology in understanding the role of soil microbial diversity for plant productivity and coexistence. Recent research has shown increases in species richness of mutualistic soil fungi, the arbuscular mycorrhizal fungi (AMF), to be related to increases in aboveground productivity of plant communities. However, the impact of AMF richness on plant-plant interactions has not been determined. Moreover, it is unknown whether species-rich AMF communities can act as insurance to maintain productivity in a fluctuating environment (e.g., upon changing soil conditions). We tested the impact of four different AMF taxa and of AMF diversity (no AMF, single AMF taxa, and all four together) on competitive interactions between the legume Trifolium pratense and the grass Lolium multiflorum grown under two different soil conditions of low and high sand content. We hypothesized that more diverse mutualistic interactions (e.g., when four AMF taxa are present) can ease competitive effects between plants, increase plant growth, and maintain plant productivity across different soil environments. We used quantitative PCR to verify that AMF taxa inoculated at the beginning of the experiment were still present at the end. The presence of AMF reduced the competitive inequality between the two plant species by reducing the growth suppression of the legume by the grass. High AMF richness enhanced the combined biomass production of the two plant species and the yield of the legume, particularly in the more productive soil with low sand content. In the less productive (high sand content) soil, the single most effective AMF had an equally beneficial effect on plant productivity as the mixture of four AMF. Since contributions of single AMF to plant productivity varied between both soils, higher AMF richness would be required to maintain plant productivity in heterogeneous environments. Overall this work shows that AMF diversity promotes plant productivity and that AMF diversity can act as insurance to sustain plant productivity under changing environmental conditions.     

Plant-soil feedback: testing the generality with the same grasses in serpentine and prairie soils

Plants can alter soil properties in ways that feed back to affect plant performance. The extent that plant-soil feedback affects co-occurring plant species differentially will determine its impact on plant community structure. Whether feedback operates consistently across similar plant communities is little studied. Here, the same grasses from two eastern U.S. serpentine grasslands and two midwestern tallgrass prairie remnants were examined for plant-soil feedback in parallel greenhouse experiments. Native soils were homogenized and cultured (trained) for a year with each of the four grasses. Feedback was evaluated by examining biomass variation in a second generation of (tester) plants grown in the trained soils. Biomass was lower in soils trained by conspecifics compared to soils trained by heterospecifics in seven of 15 possible cases; biomass was greater in conspecific soils in one other. Sorghastrum nutans exhibited lower biomass in conspecific soils for all four grasslands, so feedback may be characteristic of this species. Three cases from the Hayden prairie site were explained by trainer species having similar effects across all tester species so the relative performance of the different species was little affected; plants were generally larger in soils trained by Andropogon gerardii and smaller in soils trained by S. nutans. Differences among sites in the incidence of feedback were independent of serpentine or prairie soils. To explore the causes of the feedback, several soil factors were measured as a function of trainer species: nutrients and pH, arbuscular mycorrhizal (AM) spore communities, root colonization by AM fungi and putative pathogens, and functional diversity in bacterial communities as indicated by carbon substrate utilization. Only variation in nutrients was consistent with any patterns of feedback, and this could explain the greater biomass in soils trained by A. gerardii at Hayden. Feedback at Nottingham (one of the serpentine sites) differed, most notably for A. gerardii, from that of similar past studies that used different experimental protocols. To understand the consequences of feedback for plant community structure, it is important to consider how multiple species respond to the same plant-induced soil variation as well as differences in the feedback detected between greenhouse and field settings.     

Specificity between Neotropical tree seedlings and their fungal mutualists leads to plant-soil feedback

A growing body of evidence obtained largely from temperate grassland studies suggests that feedbacks occurring between plants and their associated soil biota are important to plant community assemblage. However, few studies have examined the importance of soil organisms in driving plant-soil feedbacks in forested systems. In a tropical forest in central Panama, we examined whether interactions between tree seedlings and their associated arbuscular mycorrhizal fungi (AMF) lead to plant-soil feedback. Specifically, do tropical seedlings modify their own AMF communities in a manner that either favors or inhibits the next cohort of conspecific seedlings (i.e., positive or negative feedback, respectively)? Seedlings of two shade-tolerant tree species (Eugenia nesiotica, Virola surinamensis) and two pioneer tree species (Luehea seemannii, Apeiba aspera) were grown in pots containing identical AMF communities composed of equal amounts of inoculum of six co-occurring AMF species. The different AMF-host combinations were all exposed to two light levels. Under low light (2% PAR), only two of the six AMF species sporulated, and we found that host identity did not influence composition of AMF spore communities. However, relative abundances of three of the four AMF species that produced spores were influenced by host identity when grown under high light (20% PAR). Furthermore, spores of one of the AMF species, Glomus geosporum, were common in soils of Luehea and Eugenia but absent in soils of Apeiba and Virola. We then conducted a reciprocal experiment to test whether AMF communities previously modified by Luehea and Apeiba differentially affected the growth of conspecific and heterospecific seedlings. Luehea seedling growth did not differ between soils containing AMF communities modified by Luehea and Apeiba. However, Apeiba seedlings were significantly larger when grown with Apeiba-modified AMF communities, as compared to Apeiba seedlings grown with Luehea-modifed AMF communities. Our experiments suggest that interactions between tropical trees and their associated AMF are species-specific and that these interactions may shape both tree and AMF communities through plant-soil feedback.     

Effects of Arbuscular mycorrhizal fungi isolated from white clovers (Trifolium repens L) on soil bacteria and fungi

White clover potted experiments were performed to investigate the effects of seven indigenous arbuscular mycorrhizal fungi (AMF) communities isolated from different test plots subjected to long-term fertilisation on soil enzyme activities, number of soil bacteria and fungi. The results showed that the inoculation of arbuscular mycorrhizal fungi communities increased the mycorrhizal infection rate of the plants and promoted the growth of plants. The Mnp treatment was most effective. The shoot biomass, root biomass, potassium and nitrogen uptake of the white clover in Mnp treatment group were increased by 61.54%, 84.00%, 62.50% and 46.71% respectively, compared with those in non-inoculation treatment. The inoculation of AMF communities had little effect on the number of bacteria in the soil, but significantly increased the number of soil fungi. Mnk treatment group had the highest number of fungi in the soil, which was 9.91 times that of the non-inoculation treatment group. The catalase and dehydrogenase activities were both significantly improved in Mnp treatment by 28.12% and 205.38% respectively, compared with those of the control treatment (-M). The urease, invertase and cellulase activities reached the highest levels in the Mck treatment; they were increased by 142.79%, 41.17% and 77.62% respectively, compared with those of the control treatment. Pearson correlation analysis showed that the soil enzyme activity was not correlated with the mycorrhizal infection rate, but correlated with the spore number of the AMF community. The impact of AMF community on soil quality is important for us to understand the function of the ecosystems. Relevant study provides important guidance for maintaining the balance of the soil-plant system and the development of sustainable agriculture.     

Numerical modelling of biostabilisation for a coal mine overburden dump slope

An integrated study on biological stabilisation of the dump slope has indicated that biological reclamation should be considered for long term stability of dump. The grasses have good soil binding capacity and help to control soil erosion and improve dump stability. Native grasses viz. Bamboo (Dendrocalmus strictus) and Kashi (Saccharum spontaneum) are the important constituents of grass species which can stabilise the dump slopes. Field observation of root development of these grass species has indicated that the roots can proliferate upto 0.5 m depth on a coal mine overburden dump after three years of vegetation. From the numerical modelling it has been analysed that roots of these grasses which significantly enhance the factor of safety of dump slope from 1.2 to 1.4 have a positive role in maintaining long term stability.     

Bromus tectorum invasion alters nitrogen dynamics in an undisturbed arid grassland ecosystem

The nonnative annual grass Bromus tectorum has successfully replaced native vegetation in many arid and semiarid ecosystems. Initial introductions accompanied grazing and agriculture, making it difficult to separate the effects of invasion from physical disturbance. This study examined N dynamics in two recently invaded, undisturbed vegetation associations (C3 and C4). The response of these communities was compared to an invaded/ disturbed grassland. The invaded/disturbed communities had higher surface NH4+ input in spring, whereas there were no differences for surface input of NO3-. Soil inorganic N was dominated by NH4+, but invaded sites had greater subsurface soil NO3-. Invaded sites had greater total soil N at the surface four years post-invasion in undisturbed communities, but total N was lower in the invaded/disturbed communities. Soil delta15N increased with depth in the noninvaded and recently invaded communities, whereas the invaded/disturbed communities exhibited the opposite pattern. Enriched foliar delta15N values suggest that Bromus assimilated subsurface NO3-, whereas the native grasses were restricted to surface N. A Rayleigh distillation model accurately described decomposition patterns in the noninvaded communities where soil N loss is accompanied by increasing soil delta15N; however, the invaded/ disturbed communities exhibited the opposite pattern, suggesting redistribution of N within the soil profile. This study suggests that invasion has altered the mechanisms driving nitrogen dynamics. Bromus litter decomposition and soil NO3- concentrations were greater in the invaded communities during periods of ample precipitation, and NO3- leached from the surface litter, where it was assimilated by Bromus. The primary source of N input in these communities is a biological soil crust that is removed with disturbance, and the lack of N input by the biological soil crust did not balance N loss, resulting in reduced total N in the invaded/disturbed communities. Bromus produced a positive feedback loop by leaching NO3- from decomposing Bromus litter to subsurface soil layers, accessing that deepsoil N pool with deep roots and returning that N to the surface as biomass and subsequent litter. Lack of new inputs combined with continued loss will result in lower total soil N, evidenced by the lower total soil N in the invaded/disturbed communities.