Nitrogen-fixing bacteria alleviates competition between arbuscular mycorrhizal fungi and Solidago canadensis for nutrients under nitrogen limitation

丛枝菌根真菌(AMF)能与大多数陆生植物的根系形成共生体, 有助于宿主植物吸收养分。但营养胁迫下, 根系微生物对AMF与宿主植物间关系的影响少见报道。该研究假设: 在营养极度匮乏(如氮胁迫)环境下, AMF与宿主植物可能产生营养竞争, 而固氮菌的介入能够缓解两者对营养的竞争关系。为了验证这一假设, 该文探究了加拿大一枝黄花(Solidago canadensis)生长受限的氮浓度, 并在氮受限条件下检验了AMF、加拿大一枝黄花及固氮菌三者间的关系。结果表明: 低氮处理明显抑制了加拿大一枝黄花的地上生物量和总生物量, 尤其以0.025 mmol·L^-1 N的氨态氮对加拿大一枝黄花的负影响更甚。在此氮浓度下, 单独添加AMF总体上都进一步抑制了加拿大一枝黄花的生长, 而固氮菌的添加在一定程度上提高了氮受限条件下AMF对宿主的根部侵染率及宿主植物生物量。这表明固氮菌能够缓和氮受限条件下AMF和加拿大一枝黄花间的营养竞争关系。研究结果加深了对外来植物在极度营养胁迫环境下与多种微生物互作的入侵机制的理解。     

固氮菌缓解氮限制环境中丛枝菌根真菌对加拿大一枝黄花的营养竞争

丛枝菌根真菌(AMF)能与大多数陆生植物的根系形成共生体, 有助于宿主植物吸收养分。但营养胁迫下, 根系微生物对AMF与宿主植物间关系的影响少见报道。该研究假设: 在营养极度匮乏(如氮胁迫)环境下, AMF与宿主植物可能产生营养竞争, 而固氮菌的介入能够缓解两者对营养的竞争关系。为了验证这一假设, 该文探究了加拿大一枝黄花(Solidago canadensis)生长受限的氮浓度, 并在氮受限条件下检验了AMF、加拿大一枝黄花及固氮菌三者间的关系。结果表明: 低氮处理明显抑制了加拿大一枝黄花的地上生物量和总生物量, 尤其以0.025 mmol·L^-1 N的氨态氮对加拿大一枝黄花的负影响更甚。在此氮浓度下, 单独添加AMF总体上都进一步抑制了加拿大一枝黄花的生长, 而固氮菌的添加在一定程度上提高了氮受限条件下AMF对宿主的根部侵染率及宿主植物生物量。这表明固氮菌能够缓和氮受限条件下AMF和加拿大一枝黄花间的营养竞争关系。研究结果加深了对外来植物在极度营养胁迫环境下与多种微生物互作的入侵机制的理解。     

Differential modulation of host plant delta13C and delta18O by native and nonnative arbuscular mycorrhizal fungi in a semiarid environment

Native, drought-adapted arbuscular mycorrhizal fungi (AMF) often improve host-plant performance to a greater extent than nonnative AMF in dry environments. However, little is known about the physiological basis for this differential plant response. Seedlings of Olea europaea and Rhamnus lycioides were inoculated with either a mixture of eight native Glomus species or with the nonnative Glomus claroideum before field transplanting in a semiarid area. Inoculation with native AMF produced the greatest improvement in nutrient and water status as well as in long-term growth for both Olea and Rhamnus. Foliar delta18O measurements indicated that native AMF enhanced stomatal conductance to a greater extent than nonnative AMF in Olea and Rhamnus.delta13C data showed that intrinsic water-use efficiency in Olea was differentially stimulated by native AMF compared with nonnative AMF. Our results suggest that modulation of leaf gas exchange by native, drought-adapted AMF is critical to the long-term performance of host plants in semiarid environments. delta18O can provide a time-integrated measure of the effect of mycorrhizal infection on host-plant water relations.     

An arbuscular mycorrhizal fungus significantly modifies the soil bacterial community and nitrogen cycling during litter decomposition

Arbuscular mycorrhizal fungi (AMF) perform an important ecosystem service by improving plant nutrient capture from soil, yet little is known about how AMF influence soil microbial communities during nutrient uptake. We tested whether an AMF modifies the soil microbial community and nitrogen cycling during litter decomposition. A two‐chamber microcosm system was employed to create a root‐free soil environment to control AMF access to ¹³C‐ and ¹⁵N‐labelled root litter. Using a 16S rRNA gene microarray, we documented that approximately 10% of the bacterial community responded to the AMF, Glomus hoi. Taxa from the Firmicutes responded positively to AMF, while taxa from the Actinobacteria and Comamonadaceae responded negatively to AMF. Phylogenetic analyses indicate that AMF may influence bacterial community assembly processes. Using nanometre‐scale secondary ion mass spectrometry (NanoSIMS) we visualized the location of AMF‐transported ¹³C and ¹⁵N in plant roots. Bulk isotope ratio mass spectrometry revealed that the AMF exported 4.9% of the litter ¹⁵N to the host plant (Plantago lanceolata L.), and litter‐derived ¹⁵N was preferentially exported relative to litter‐derived ¹³C. Our results suggest that the AMF primarily took up N in the inorganic form, and N export is one mechanism by which AMF could modify the soil microbial community and decomposition processes.     

Current developments in arbuscular mycorrhizal fungi research and its role in salinity stress alleviation: a biotechnological perspective

Arbuscular mycorrhizal fungi (AMF) form widespread symbiotic associations with 80% of known land plants. They play a major role in plant nutrition, growth, water absorption, nutrient cycling and protection from pathogens, and as a result, contribute to ecosystem processes. Salinity stress conditions undoubtedly limit plant productivity and, therefore, the role of AMF as a biological tool for improving plant salt stress tolerance, is gaining economic importance worldwide. However, this approach requires a better understanding of how plants and AMF intimately interact with each other in saline environments and how this interaction leads to physiological changes in plants. This knowledge is important to develop sustainable strategies for successful utilization of AMF to improve plant health under a variety of stress conditions. Recent advances in the field of molecular biology, “omics” technology and advanced microscopy can provide new insight about these mechanisms of interaction between AMF and plants, as well as other microbes. This review mainly discusses the effect of salinity on AMF and plants, and role of AMF in alleviation of salinity stress including insight on methods for AMF identification. The focus remains on latest advancements in mycorrhizal research that can potentially offer an integrative understanding of the role of AMF in salinity tolerance and sustainable crop production.     

AM真菌物种多样性:生态功能、影响因素及维持机制

AM真菌物种多样性是土壤生态系统生物多样性的重要组分之一。尽管对AM真菌多样性已有多年研究,但是,已有研究绝大多数仅停留在对AM真菌群落种属解析层面上,对AM真菌物种多样性生态功能及维持机制方面的认识较浅。从生态功能、影响因素及维持机制3个方面系统地综述了近年来AM真菌多样性领域的研究进展。认为AM真菌多样性对植物群落生产力的调控机制及结合理论与实践解析AM真菌多样性维持机制是该领域未来的重点研究方向。     

Nitrogen and phosphorus additions impact arbuscular mycorrhizal abundance and molecular diversity in a tropical montane forest

Increased nitrogen (N) depositions expected in the future endanger the diversity and stability of ecosystems primarily limited by N, but also often co‐limited by other nutrients like phosphorus (P). In this context a nutrient manipulation experiment (NUMEX) was set up in a tropical montane rainforest in southern Ecuador, an area identified as biodiversity hotspot. We examined impacts of elevated N and P availability on arbuscular mycorrhizal fungi (AMF), a group of obligate biotrophic plant symbionts with an important role in soil nutrient cycles. We tested the hypothesis that increased nutrient availability will reduce AMF abundance, reduce species richness and shift the AMF community toward lineages previously shown to be favored by fertilized conditions. NUMEX was designed as a full factorial randomized block design. Soil cores were taken after 2 years of nutrient additions in plots located at 2000 m above sea level. Roots were extracted and intraradical AMF abundance determined microscopically; the AMF community was analyzed by 454‐pyrosequencing targeting the large subunit rDNA. We identified 74 operational taxonomic units (OTUs) with a large proportion of Diversisporales. N additions provoked a significant decrease in intraradical abundance, whereas AMF richness was reduced significantly by N and P additions, with the strongest effect in the combined treatment (39% fewer OTUs), mainly influencing rare species. We identified a differential effect on phylogenetic groups, with Diversisporales richness mainly reduced by N additions in contrast to Glomerales highly significantly affected solely by P. Regarding AMF community structure, we observed a compositional shift when analyzing presence/absence data following P additions. In conclusion, N and P additions in this ecosystem affect AMF abundance, but especially AMF species richness; these changes might influence plant community composition and productivity and by that various ecosystem processes.     

Genetic diversity and host plant preferences revealed by simple sequence repeat and mitochondrial markers in a population of the arbuscular mycorrhizal fungus Glomus intraradices

Arbuscular mycorrhizal fungi (AMF) are important symbionts of plants that improve plant nutrient acquisition and promote plant diversity. Although within-species genetic differences among AMF have been shown to differentially affect plant growth, very little is actually known about the degree of genetic diversity in AMF populations. This is largely because of difficulties in isolation and cultivation of the fungi in a clean system allowing reliable genotyping to be performed. A population of the arbuscular mycorrhizal fungus Glomus intraradices growing in an in vitro cultivation system was studied using newly developed simple sequence repeat (SSR), nuclear gene intron and mitochondrial ribosomal gene intron markers. The markers revealed a strong differentiation at the nuclear and mitochondrial level among isolates. Genotypes were nonrandomly distributed among four plots showing genetic subdivisions in the field. Meanwhile, identical genotypes were found in geographically distant locations. AMF genotypes showed significant preferences to different host plant species (Glycine max, Helianthus annuus and Allium porrum) used before the fungal in vitro culture establishment. Host plants in a field could provide a heterogeneous environment favouring certain genotypes. Such preferences may partly explain within-population patterns of genetic diversity.     

丛枝菌根真菌与氮添加对不同根形态基因型水稻氮吸收的影响

植物主要依赖自身根系从土壤中获取矿质养分; 具有不同根形态的植物对于养分的吸收能力存在差异。丛枝菌根真菌(AMF)能与陆地植物根系形成共生关系, 帮助植物吸收矿质养分。但是, AMF对于植物根系养分吸收的促进效应是否会受根形态的影响还鲜有研究。该研究选取4种不同根形态基因型水稻(根毛缺陷突变体rhl1、侧根缺陷突变体iaa11、不定根缺失突变体arl1和野生型Kas)为研究对象, 设置2种施氮水平处理(低氮: 20 mg·kg^-1氨氮; 高氮: 100 mg·kg^-1氨氮), 利用稳定同位素¹⁵N示踪标记技术, 探究AMF和氮添加对不同根形态植物氮吸收的影响。研究结果发现, 相比低氮处理, 高氮处理下, rhl1、Kas、iaa11与arl1的茎叶¹⁵N浓度分别提高了60%、72%、128%与118%, 说明氮添加显著促进了水稻氮吸收, 且iaa11与arl1对氮添加的响应更强烈。在低氮水平下, AMF对rhl1、Kas、iaa11与arl1氮吸收的平均效应值分别为17%、31%、42%、51%, 表明AMF对于植物氮吸收的促进效应受根形态影响, iaa11与arl1对AMF的响应明显高于Kas与rhl1; 相较于低氮水平, 高氮水平下AMF对于不同根形态水稻氮吸收的促进效应都会显著降低, 表明氮添加削弱了AMF对植物氮吸收的促进效应。该研究阐明了4种不同根形态基因型水稻氮养分吸收存在显著差异, 其中氮吸收能力较弱的基因型水稻对AMF的响应更强, 该结果补充了植物与AMF在养分吸收上存在功能互补的控制实验证据。     

Interaction between Earthworms and Arbuscular Mycorrhizal Fungi in Plants: A Review

Different kinds of soil animals and microorganisms inhabit the plant rhizosphere, which function closely to plant roots. Of them, arbuscular mycorrhizal fungi (AMF) and earthworms play a critical role in sustaining the soil-plant health. Earthworms and AMF belong to the soil community and are soil beneficial organisms at different trophic levels. Both of them improve soil fertility and structural development, collectively promoting plant growth and nutrient acquisition capacity. Earthworm activities redistribute mycorrhizal fungi spores and give diversified effects on root mycorrhizal fungal colonization. Dual inoculation with both earthworms and AMF strongly magnifies the response on plant growth through increased soil enzyme activities and changes in soil nutrient availability, collectively mitigating the negative effects of heavy metal pollution in plants and soils. This thus enhances phytoremediation and plant disease resistance. This review simply outlines the effects of earthworms and AMF on the soil-plant relationship. The effects of earthworms on root AMF colonization and activities are also analyzed. This paper also summarizes the interaction between earthworms and AMF on plants along with suggested future research.     

Differential influence of native and introduced arbuscular mycorrhizal fungi on growth of dominant and subordinate plants

The impact of arbuscular mycorrhizal fungi (AMF) on plant ecosystems has been intensively reported. In this research, we explored the difference between native and introduced AMF in promoting the growth of dominant and subordinate plant species. In glasshouse experiments, dominants and subordinates from subtropical grasslands were colonized by native AMF or introduced AMF, Glomus versiforme. The biomass revealed that mycorrhizal dependencies (MD) on the native AMF of the dominants were much higher than those of the subordinates, while MD on the introduced AMF changed following the replacement of native AMF with introduced AMF. A close relationship between biomass promotion and increase in phosphorus uptake was observed, indicating the important role of AMF-enhanced nutrient acquisition by roots. Our results show that plant community structures are partly determined by MD on native AMF, and could be modified by introducing exogenous AMF species.     

A knapweed biological control perspective

Diffuse and spotted knapweed (Centaurea diffusa Lam. and C. stoebe micranthos (Gugler) Hayek) are Eurasian plants that devastate dry and mesic North American grasslands. They have a mutualistic association with arbuscular mycorrhizal fungal (AMF) phylotypes with hyphal links to nearby plants and a nutrient flux to the strongest sink, usually knapweed. They displace many AMF beneficial to grass and affect knapweed nutrient allocation, biology, knapweed insects and probably root necrosis and emergence of ant buried seed. AMF determined nutrient root or shoot allocation determines nutrient shoot and root allocation and the benefit to root or seed-head insect species and whether C. diffusa is an annual–biannual or a semelparous perennial needing 5 or more years to flower. Both knapweeds do well without its AMF phylotypes without competition in fertile soil. In grass in Eurasia, they have a community of seven seed-head species segregated by head development stage. Prolonged seed dormancy buffered knapweed decline that resulted in release of a surfeit seed-head species. The presence of an eliasome on the seed and vigorous seedling clumps suggests burial by myrmecochorous ants with AMF supplied carbon supporting their growth. The root species community is segregated by habitat, climate, root part, and size. With larval induced compensatory growth and AMF nutrient sharing, the growth of plants with and without a larva was the same. On feeding completion, a nutrient out flux from the attacked plants reduced growth; but without killing. This needs a dual species or a repeated single species attack. Root species packing increases knapweed utilization; but the four approved species are insufficient for maximum utilization. Two additions are suggested. The aim of the paper is to provide enough understanding of the AMF and its plant and insect interactions to facilitate knapweed biological control and avoid past mistakes.     

Soil organisms shape the competition between grassland plant species

Decomposers and arbuscular mycorrhizal fungi (AMF) both determine plant nutrition; however, little is known about their interactive effects on plant communities. We set up a greenhouse experiment to study effects of plant competition (one- and two-species treatments), Collembola (Heteromurus nitidus and Protaphorura armata), and AMF (Glomus intraradices) on the performance (above- and belowground productivity and nutrient uptake) of three grassland plant species (Lolium perenne, Trifolium pratense, and Plantago lanceolata) belonging to three dominant plant functional groups (grasses, legumes, and herbs). Generally, L. perenne benefited from being released from intraspecific competition in the presence of T. pratense and P. lanceolata. However, the presence of AMF increased the competitive strength of P. lanceolata and T. pratense against L. perenne and also modified the effects of Collembola on plant productivity. The colonization of roots by AMF was reduced in treatments with two plant species suggesting that plant infection by AMF was modified by interspecific plant interactions. Collembola did not affect total colonization of roots by AMF, but increased the number of mycorrhizal vesicles in P. lanceolata. AMF and Collembola both enhanced the amount of N and P in plant shoot tissue, but impacts of Collembola were less pronounced in the presence of AMF. Overall, the results suggest that, by differentially affecting the nutrient acquisition and performance of plant species, AMF and Collembola interactively modify plant competition and shape the composition of grassland plant communities. The results suggest that mechanisms shaping plant community composition can only be understood when complex belowground interactions are considered.     

典型农田土壤中丛枝菌根真菌-根际细菌互作及与氮磷利用的关系

在农业生态系统中,土壤微生物是土壤-作物系统养分循环的重要驱动力,其中丛枝菌根真菌(Arbuscularmycorrhizalfungi,AMF)能够促进作物对养分的吸收,适应逆境胁迫。【目的】进一步揭示AMF和根际细菌群落的跨界网络互作,挖掘与作物氮磷利用显著相关的关键微生物类群,揭示关键类群的生态网络特征。【方法】利用Illumina测序技术对3种典型农田旱地土壤(黑土、潮土和红壤)中AMF和根际细菌群落结构进行分析;构建互作网络并利用偏冗余分析、相关性分析探究了与氮磷利用相关的潜在关键类群。【结果】3种土壤中AMF与根际细菌均以正相互作用为主。不同土壤中AMF与根际细菌互作关系差异明显,在红壤中跨界互作最为密切,其中球囊霉属真菌(Glomus)与根际细菌中的放线菌(Actinobacteria)和变形菌(Proteobacteria)之间的交互作用最多。而在黑土中主要体现为根际细菌的界内互作。与氮磷利用率显著相关的关键微生物类群主要属于球囊霉属真菌、放线菌和α变形菌。【结论】典型旱地土壤中AMF与根际细菌的正相互关系对作物氮磷利用有潜在促进作用,关键类群在有机质和养分贫乏的红壤中可能起到更重要的作用。     

Silicon influences growth and mycorrhizal responsiveness in strawberry plants

Effect of silicon (Si) on the response of strawberry (Fragaria?×?ananassa var. Parus) plants to arbuscular mycorrhizal fungus (AMF) was studied under growth chamber conditions. Plants were grown in perlite irrigated with nutrient solution without (??Si) or with (+?Si) 3 mmol L^?1 Si (~?84 mg L^?1 Si as Na₂SiO₃) in the absence (??AMF) or presence (+?AMF) of fungus. Dry matter production, root colonization rate, photosynthesis rate and water relation parameters were all improved by both Si and AMF, and the highest amounts were achieved by +?Si?+?AMF treatment. Mycorrhizal effectiveness increased by Si treatment associated with higher Si concentration in the +?AMF plants. Leaf concentrations of total soluble and cell wall-bound phenolics were increased by Si accompanied by the enhanced activity of phenylalanine ammonia lyase, but not polyphenol oxidase. Profile of phenolics compound revealed that gallic acid, caffeic acid, epicatechin, chlorogenic acid, ellagic acid and kaempferol increased by both Si and AMF treatments, while p-coumaric acid decreased. In addition to vegetative growth, both treatments improved fruit yield and its quality parameters. Our results showed that Si and AMF acted in a synergistic manner and improved growth and biochemical parameters in strawberry plants. However, the mechanism for Si-mediated increase of mycorrhizal effectiveness is not known, thereby needing further elucidation.     

丛枝菌根真菌和施加不同形态氮肥对杉木幼苗养分吸收的影响

为了解丛枝菌根真菌(AMF)和不同形态氮对杉木(Cunninghamia lanceolata)生长和养分吸收的影响,以1 a生杉木幼苗接种摩西球囊霉(Glomus mosseae)和添加不同形态氮(NH₄⁺-N和NO₃^–-N),对其养分元素和生长状况的变化进行研究。结果表明,AMF显著提高了杉木的苗高和生物量,促进了杉木对N、P、K、Ca、Mg、Fe和Na的吸收,AMF对微量元素Fe、Na的促进作用总体上要强于大量元素K、Ca。与NO₃^–-N相比,AMF显著提高了NH₄⁺-N处理杉木的生物量、总C和N、Ca、Mg、Mn含量,而且这种显著性在叶中普遍高于根和茎。接种AMF可以促进杉木幼苗的生长和对养分元素的吸收,且添加NH₄⁺-N处理的促进作用要强于NO₃^–-N。     

Arbuscular mycorrhizal fungi alter plant community composition along a grazing gradient in Inner Mongolia Steppe

Arbuscular mycorrhizal fungi (AMF) have a significant influence on plant productivity and diversity in non-grazing grassland. However, the interactive effects between grazing intensity and AMF on plant community composition in natural grassland communities are not well known. We conducted a field experiment that manipulated AMF colonization and grazing intensity to study the impact of AMF suppression on plant community composition and nutrient status over 2 years (2015–2016) with contrasting rainfall levels. We found that AMF root colonization was significantly reduced by the application of the fungicide benomyl as a soil drench. Grazing intensity regulated plant community composition and aboveground biomass mainly by reducing the growth of Leymus chinensis over 2 years. AMF suppression increased the growth of Chenopodium glaucum, but it did not alter other plant species across all grazing intensities. The effects of AMF suppression on plant community composition changed along a grazing gradient considerably between years: AMF suppression increased the biomass of C. glaucum across all grazing intensities in 2015, but slightly increased it in 2016. Interactions between AMF suppression and grazing intensity altered the phosphorus concentration of Stipa grandis and Cleistogenes squarrosa in 2015 but not in 2016. AMF suppression decreased the shoot phosphorus content of L. chinensis but increased that of C. glaucum across all grazing intensities. Our results indicate that grazing intensity substantially alters aboveground community biomass and affects growth of dominant species; AMF by itself have limited effects on plant communities along a grazing gradient in typical steppe.     

Mycorrhizal‐mediated nitrogen acquisition in switchgrass under elevated temperatures and N enrichment

Arbuscular mycorrhizal fungi (AMF) can perform key roles in ecosystem functioning through improving host nutrient acquisition. Nitrogen (N) is an essential nutrient for plant growth, however, anthropogenic N loading (e.g. crop fertilization and deposition from combustion sources) is increasing so that N now threatens ecosystem sustainability around the world by causing terrestrial and aquatic eutrophication and acidification. It is important to better understand the capacity of AMF to directly uptake N from soils and transfer it to host plants because this process may increase N recycling and retention within ecosystems. In addition to understanding the role of AMF in the N cycle in the present day it is important to understand how AMF function may change as global change proceeds. Currently the net effects of N enrichment and elevated temperature predicted with global change on AMF are unknown. In this study, we examined the effects of N enrichment by simulated N‐deposition loading, elevated temperatures expected by future global changes and their interactions on growth and AMF‐mediated N acquisition of switchgrass (Panicum virgatum var. Alamo), an important species for biofuel production. Switchgrass plants were grown in microcosm units that divided mycorrhizal roots from AMF hyphae and organic residues enriched with ¹⁵N by compartments separated by an air gap to reduce N diffusion. While AMF did not enhance switchgrass biomass, mycorrhizas significantly increased ¹⁵N in shoots and total shoot N. Neither N enrichment nor elevated temperatures influenced this mycorrhizal‐mediated N uptake and transfer. Results from this study can aid in developing sustainable bioethanol and switchgrass production practices that are less reliant on synthetic fertilizers and more dependent on internal N recycling from AMF.     

Effects of nutrient and soil moisture on competition between shape Carex stricta,shape Phalaris arundinacea,and shape Typha latifolia

We investigated the importance of nutrients, soil moisture, arbuscular mycorrhizal fungi (AMF), and interspecific competition levels on the biomass allocation patterns of three wetland perennial plant species, Carex stricta Lam., Phalaris arundinacea L., and Typha latifolia L. A factorial experiment was conducted with high-low nutrient levels, high-low soil moisture levels, and with and without AMF inoculation. Under the experimental conditions, plant inoculation by AMF was too low to create a treatment and the AMF treatment was dropped from the total analysis. P. arundinacea and T. latifolia biomass were 73% and 77% higher, respectively, in the high nutrient treatment compared to the low nutrient treatment. Biomass allocation between shoots and roots remained relatively constant between environmental treatments, although shoot:root ratios of P. arundinacea declined in the low nutrient treatment. For C. stricta, the high nutrient and soil moisture treatments resulted in an increase in biomass of 50% and 15%, respectively. Shoot:root ratios were nearly constant among all environmental conditions. Biomass of T. latifolia and C. stricta was greatly decreased when grown with P. arundinacea. The rapid, initial height growth of P. arundinacea produced a spreading, horizontal canopy that overshadowed the vertical leaves of T. latifolia and C. stricta throughout the study. This pattern was repeated in both high and low nutrient and soil moisture treatments. When grown with P. arundinacea, C. stricta and T. latifolia significantly increased their mean shoot height, regardless of the nutrient or soil moisture level. The results of this experiment suggest that C. stricta and T. latifolia were light limited when growing with P. arundinacea and that canopy architecture is more important for biomass allocation than the other environmental conditions tested. The results also suggest that Phalaris arundinacea is an inherently better competitor (sensu Grime 1979) than C. stricta or T. latifolia.