Endomycorrhizal fungi in nitrogen transfer from soybean to maize

Using ¹⁵N as a tracer, interspecific N-transfer was studied during the course of plant development. The use of barriers of differing permeabilities between donor and receiver plants allowed separation of the effect of mycorrhizal colonization, root or hyphal contact and interplant hyphal bridging, on ¹⁵N-transfer from soybean (Glycine max (L.) Merrill) to maize (Zea mays L.). More transfer was measured between mycorrhizal plants, but transport of ¹⁵N from the labelled host plant to Glomus versiforme (Karsten) Berch did not seem to occur at the symbiotic interface, suggesting that the fungus is independent of its host for its N-nutrition, and that the role of hyphal bridges in N-transfer between plants, is not significant. Uptake by the receiver plant of the N excreted by the donor plant root system appears to be the mechanism of N-transfer between plants. The factor most affecting ¹⁵N-transfer between plants was found to be the extent of the contact between plant root systems. The presence of the endomycorrhizal fungus in plant roots reduced ¹⁵N-loss from soybean, but at the same time, its extensive hyphal network improved the efficiency of the maize root system for the recovery of the ¹⁵N excreted by soybeans. The net result was a better conservation of the N resource within the plant system. The transfer of N between mycorrhizal plants was particularly enhanced by the death of the soybean.     

丛枝菌根根外菌丝网络形成过程中的时间效应及植物介导作用

以豆科植物紫花苜蓿为试验材料,应用三室(供体室-间隔室-受体室)培养系统,研究在供体和受体紫花苜蓿根系之间菌丝网络形成的时间效应以及间隔室中不同植物对菌丝网络建成的介导作用.第一个试验在供体和受体植物生长8、10、12、14周之后进行收获以检验菌丝网络形成的时间效应;第二个试验则在间隔室分别种植紫花苜蓿、羊草和独行菜,以考察菌根依赖性不同的植物对菌丝网络形成的介导作用.试验结果显示:(1)接种丛枝菌根真菌的供体紫花苜蓿根系能够形成良好的菌根共生,其外延菌丝可穿过尼龙网和间隔室侵染受体植物根系;植物生长8周后,在受体植物根系检测到菌根侵染,证实供体和受体植物间形成了根间菌丝网络;10周后,尽管供体室和受体室植物的侵染率已无差异,但二者的生物量和地上部磷浓度差异却加大,表现出菌丝网络对植物种内竞争影响的不对称性.(2)试验条件下,不同介导植物对受体植物的菌根侵染及生物量均无明显影响,但显著降低了供体植物生物量和地上部磷浓度;间隔室无介导植物或种植独行菜时,受体植物地上部和根系生物量显著低于供体植物,而当介导植物为紫花苜蓿和羊草时,受体和供体植物生物量无显著差异.研究表明,植物根间菌丝网络的形成受时间和介导植物的影响,同时也具有调节植物间资源分配和植物相互作用的功能.     

A comparison of phosphorus and nitrogen transfer between plants of different phosphorus status

Summary This study has two objections: (1) to compare transfers of phosphorus (³²P) with nitrogen (¹⁵N) from undefoliated and defoliated mycorrhizal P-rich plants to an adjacent mycorrhizal plant and (2) to determine whether the improved nutrient status of a plant growing with a nutrient-rich plant is due primarily to movement of nutrients from roots of its nutrient-rich neighbor (= nutrient transfer) or to reduced nutrient uptake by its nutrient-rich neighbor (=shift in competition). Two plants of Plantago lanceolata were grown in a three-pot unit in which each of their root systems were split, with part in the central shared pot and part by themselves in an outside pot. There were three treatments: (1) no added P; (2) P added in the outer pot to only plant, termed the donor plant, since it might provide P to the companion plant, acting as a receiver; and (3) as in the previous treatment but the P-fertilized donor plant was also clipped. To encourage the formation of hyphal links between roots of the different plants, transfers were determined when root length densities were high (90 to 130 cm cm^-3 soil) and when 56 to 85% of the root length was infected with vesicular-arbuscular mycorrhizae. Phosphorus fertilization enhanced P but not N movement within donor plants. Regardless of treatment, N transfer from donor to receiver plants was an order of magnitude greater than P transfer and in amounts that could potentially affect plant nutrition in very infertile soils. Phosphorus transfer was very small in any of the treatments. Although P fertilization and clipping improved P status of receiver plants, P transfer was not indicated as the main reason for the improved nutrition. A shift in competition between donor and receiver plants was likely the major factor in the shift in nutrition of the receiver plants.     

The symbiotic recapture of nitrogen from dead mycorrhizal and non-mycorrhizal roots of tomato plants

Aims

The aim was to quantify the nitrogen (N) transferred via the extra-radical mycelium of the arbuscular mycorrhizal fungus Glomus intraradices from both a dead host and a dead non-host donor root to a receiver tomato plant. The effect of a physical disruption of the soil containing donor plant roots and fungal mycelium on the effectiveness of N transfer was also examined.

Methods

The root systems of the donor (wild type tomato plants or the mycorrhiza-defective rmc mutant tomato) and the receiver plants were separated by a 30 μm mesh, penetrable by hyphae but not by the roots. Both donor genotypes produced a similar quantity of biomass and had a similar nutrient status. Two weeks after the supply of ¹⁵?N to a split-root part of donor plants, the shoots were removed to kill the plants. The quantity of N transferred from the dead roots into the receiver plants was measured after a further 2 weeks.

Results

Up to 10.6 % of donor-root ¹⁵N was recovered in the receiver plants when inoculated with the arbuscular mycorrhizal fungus (AMF). The quantity of ¹⁵N derived from the mycorrhizal wild type roots clearly exceeded that from the only weakly surface-colonised rmc roots. Hyphal length in the donor rmc root compartments was only about half that in the wild type compartments. The disruption of the soil led to a significantly increased AMF-mediated transfer of N to the receiver plants.

Conclusions

The transfer of N from dead roots can be enhanced by AMF, especially when the donor roots have been formerly colonised by AMF. The transfer can be further increased with higher hyphae length densities, and the present data also suggest that a direct link between receiver mycelium and internal fungal structures in dead roots may in addition facilitate N transfer. The mechanical disruption of soil containing dead roots may increase the subsequent availability of nutrients, thus promoting mycorrhizal N uptake. When associated with a living plant, the external mycelium of G. intraradices is readily able to re-establish itself in the soil following disruption and functions as a transfer vessel.     

Endomycorrhizal fungal species mediate 15N transfer from soybean to maize in non-fumigated soil

The effect of mycorrhizal inoculation on ¹⁵N transfer from soybean to maize was studied in fumigated and non-fumigated soil. Three Glomus species and a non-inoculated control were compared.In spite of higher levels of root colonization and more abundant hyphae associated with plants growing in fumigated soil, mycorrhizae-enhanced ¹⁵N transfer to maize was significant only in non-fumigated plots. High ¹⁵N transfer was not only associated with high mycelium density in soil but also with low soil microbial carbon, suggesting that the effect of mycorrhizal fungi on soil microbial populations may be an important factor affecting N transfer between mycorrhizal plants.     

Specificity of interplant cycling of phosphorus: The role of mycorrhizas

The cycling of nutrients from dying roots of one plant (the donor) to other intact plants (the receivers) was examined in a series of pot experiments. In each pot receiver plants formed either the same or a different type of mycorrhiza as the donor plant and was therefore respectively either capable or incapable of forming mycorrhizal hyphal links to the donor. There was a preferential transfer of ³²P from the dying roots of the vesicular-arbuscular mycorrhizal (VAM) Lolium perenne to VAM-infected trees Acer pseudoplatanus and Fraxinus excelsior compared to the ectomycorrhizal (ECM) Larix eurolepis, this despite an apparently greater competitive ability of L. eurolepis to obtain ³²P from the soil. Following the death of L. perenne roots there was also an increase in total P in the VAM tree receiver. These findings could not be explained by similarities in rooting distribution of the VAM-infected plants.In a similar study of the transfer of ³²P between heathland plants there was a preferential cycling of ³²P from one ericoid mycorrhizal Calluna vulgaris to another rather than to the VAM Molinia caerulea. In contrast, when ³²P was supplied directly to the soil, M. caerulea obtained significantly more ³²P than C. vulgaris. These results are discussed in relation to the potential role of interplant mycorrhizal links in the cycling of nutrients within partially closed cycles and the implications that this might have for species balance in plant communities.     

Decomposition of 14C-labelled lignin,holocellulose and lignocellulose by mycorrhizal fungi

Five different species of known ecto-mycorrhizal fungi: Cenococcum geophilum, Amanita muscaria, Tricholoma aurantium, Rhizopogon luteolus and Rhizopogon roseolus were studied for their ability to metabolize the major components of plant cell walls. All strains were able to decompose ¹⁴C-labelled plant lignin, ¹⁴C-lignocellulose and ¹⁴C-DHP-lignin at a rate which was lower than the one observed for the known white rot fungi Heterobasidion annosum and Sporotrichum pulverulentum. Also ¹⁴C-(U)-holocellulose was relatively less degradable for the mycorrhizal fungi than for the white rotters. On the other hand, aromatic monomers like ¹⁴C-vanillic acid were decomposed to a much higher extent by two species of mycorrhizal fungi compared to the activity observed for Heterobasidion annosum. The results of the experiments reveal that these stains of mycorrhizal fungi are well able to utilize the major components of plant material and thus can contribute to litter decomposition in the forest floor.     

Intraspecific transfer of carbon between plants linked by a common mycorrhizal network

To quantify the involvement of arbuscular mycorrhiza (AM) fungi in the intraspecific transport of carbon (C) between plants we fumigated established Festuca ovina turf for one week with air containing depleted ¹³C. This labelled current assimilate in a section of mycorrhizal or non-mycorrhizal turf. Changes in the ^13/12C ratio of adjacent, unfumigated plants, therefore, allowed the movement of C between labelled and unlabelled plants to be estimated. In mycorrhizal turves, 41% of the C exported to the roots from the leaves was transported to neighbouring plants. The most likely explanation of this is was the transport of C via a common hyphal network connecting the roots of different plants. No inter-plant transport of C was detected in non-mycorrhizal turves. There was no evidence that the C left fungal structures and entered the roots of receiver plants. Mycorrhizal colonisation increased carbon transport from leaves to root from 10% of fixed carbon when non-mycorrhizal to 36% in mycorrhizal turves. These results suggest that AM fungi impose a significantly greater C drain on host plants than was previously thought.     

Nitrogen Transfer Between Plants: A 15N Natural Abundance Study with Crop and Weed Species

An increasing amount of evidence indicates that N can be transferred between plants. Nonetheless, a number of fundamental questions remain. A series of experiments was initiated in the field to examine N transfer between N₂-fixing soybean (Glycine max [L.] Merr.) varieties and a non-nodulating soybean, and between N₂-fixing peanut (Arachis hypogaea L.) or soybean and neighboring weed species. The experiments were conducted in soils with low N fertilities and used differences in N accumulation and/or ¹⁵N natural abundance to estimate N transfer. Mixtures of N₂-fixing and non-nod soybean indicated that substantial inter-plant N transfer occurred. Amounts were variable, ranging from negligible levels to 48% of the N found in the non-nod at maturity. Transfer did not appear to strongly penalize the N₂-fixing donor plants. But, in cases where high amounts of N were transferred, N content of donors was noticeably lowered. Differences were evident in the amount of N transferred from different N₂-fixing donor genotypes. Results of experiments with N₂-fixing crops and the weed species prickly sida (Sida spinosa L.) and sicklepod (Senna obtusifolia [L.] Irwin & Barneby) also indicated substantial N transfer occurred over a 60-day period, with amounts accounting for 30–80% of the N present in the weeds. Transfer of N, however, was generally very low in weed species that are known to be non-hosts for arbuscular mycorrhizae (yellow nutsedge, Cyperus esculentus L. and Palmer amaranth, Amaranthus palmeri [S.] Watson). The results are consistent with the view that N transfer occurs primarily through mycorrhizal hyphal networks, and they reveal that N transfer may be a contributing factor to weed problems in N₂-fixing crops in low N fertility conditions.     

Effects of vesicular-arbuscular mycorrhiza on the size of the labile pool of soil phosphate

Summary Inoculation of lettuce, onion and clover with VA mycorrhizal fungus (Glomus mosseae) increased plant yields and phosphate uptake in three soils that had been depleted in phosphate. From two soils in which the labile pool of phosphate had been labelled with³²P, the specific activity of plant phosphate was the same whether the plants were mycorrhizal or non-mycorrhizal. In a third soil (Sonning) the specific activity was lower in lettuce and clover when the plants were mycorrhizal. When the experiment was repeated with the same soil under conditions that gave lower growth rates, the specific activity was the same in mycorrhizal and non-mycorrhizal plants. The lower specific activity in lettuce and clover in the first experiment is atributed to greater release of slowly exchanging phosphate (which is not in equilibrium with the added³²P), caused by the high uptake of phosphate by the mycorrhizal plants. When they occur, lower specific activities in mycorrhizal plants may therefore not necessarily indicate a solubilizing effect of the mycorrhiza on soil phosphate.     

Carbon distribution and nitrogen partitioning in a soil-plant system with barley (Hordeum vulgare L.), ryegrass (Lolium perenne) and rape (Brassica napus L.) grown in a14CO2-atmosphere

To examine the influence of plant-microorganism interactions on soil-N transformations (e.g. net mineralization, net immobilization) a pot experiment was conducted in a¹⁴C-labelled atmosphere by using different (two annuals, one perennial) plants species. It was assumed that variation in below-ground, microorganism-available C would influence N transformations in soil. Plant species were fertilized (low rate) with¹⁵N-labelled nitrogen and grown, during days 13 and 62 after germination, in a growth chamber with a¹⁴C-labelled atmosphere. Nitrification was inhibited by using nitrapyrin (N-Serve). During the chamber period, shoots were harvested, and associated roots and soil were collected on two sampling occasionm, e.g. after 4 and 7 weeks in the growth chamber.The distribution of net (%) assimilated¹⁴C was significantly affected by both plant and time factors, and there was a significant plant × time interaction. There were significant differences between plants in all plant-soil compartments examined as well as in the degree of the plant × time interaction.Differences in the¹⁴C distribution between plants were due to both interspecific and developmental variation. In general, when comparing¹⁵N and¹⁴C quantities between species, many of the differences found between plants can be explained by the differences determined in the weight of shoot or root parts. Despite the fact that amounts of C released were greater in ryegrass than in the other plant-treatments no unequivocal evidence was found to show that the effects of plant-microorganism interactions on soil-N mineralization were greater under ryegrass. Possible mechanisms accounting for the partitioning of N found among plant biomass, soil biomass and soil residues are discussed.     

Is nitrogen transfer among plants enhanced by contrasting nutrient‐acquisition strategies?

Nitrogen (N) transfer among plants has been found where at least one plant can fix N₂. In nutrient‐poor soils, where plants with contrasting nutrient‐acquisition strategies (without N₂ fixation) co‐occur, it is unclear if N transfer exists and what promotes it. A novel multi‐species microcosm pot experiment was conducted to quantify N transfer between arbuscular mycorrhizal (AM), ectomycorrhizal (EM), dual AM/EM, and non‐mycorrhizal cluster‐rooted plants in nutrient‐poor soils with mycorrhizal mesh barriers. We foliar‐fed plants with a K¹⁵NO₃ solution to quantify one‐way N transfer from ‘donor’ to ‘receiver’ plants. We also quantified mycorrhizal colonization and root intermingling. Transfer of N between plants with contrasting nutrient‐acquisition strategies occurred at both low and high soil nutrient levels with or without root intermingling. The magnitude of N transfer was relatively high (representing 4% of donor plant N) given the lack of N₂ fixation. Receiver plants forming ectomycorrhizas or cluster roots were more enriched compared with AM‐only plants. We demonstrate N transfer between plants of contrasting nutrient‐acquisition strategies, and a preferential enrichment of cluster‐rooted and EM plants compared with AM plants. Nutrient exchanges among plants are potentially important in promoting plant coexistence in nutrient‐poor soils.     

The effect of season on VA mycorrhiza of the almond tree and of phosphate fertilization and species of endophyte on its mycorrhizal dependency

Summary Prunus dulcis (Miller), a tree which is able to develop in low fertility soils, forms VA mycorrhiza. Under glasshouse conditions the growth and P concentration in the leaf tissue of non-mycorrhizal plants, given a customary agronomic dose of P-fertilizer, were lower than those of mycorrhizal plants. The relative mycorrhizal dependency¹⁴ values of the almond tree were higher when a mixture of locally isolated (mainlyGlomus fasciculatus) was used as inoculum. These indigenous endophytes were more tolerant of added fertilizers thanGlomus mosseae taken from the pot-culture collection.The amount of VA infection and the number of Endogonaceae spores in the rhizosphere of almond trees growing in the field steadily increase from winter (the flowering season of this crop) until summer or early autumn.     

Interspecific nutrient transfer in a tallgrass prairie plant community

Interplant nutrient transfer may be an important ecological process in grasslands, and may significantly influence plant neighborhood interactions. We investigated the potential for phosphorus transfer between the dominant grass Andropogon gerardii and several neighboring plant species in tallgrass prairie via a field ³²PO₄ labelling experiment. The mean amount of ³²P received from donor shoots differed significantly among neighboring species and decreased with increasing distance from the donor. In general, forbs and cool-season C₃ grasses received more labelled ³²P than warm-season C₄ grasses. Phosphorus transfer occurred over distances up to 0.5 m. The effects of species and distance on movement of phosphorus changed with increasing time after labelling. The relative mass of receiver and donor shoots did not affect amounts of ³²P transfer. A benomyl fungicide treatment, applied to suppress mycorrhizal activity, likely did not affect existing vegetative hyphae and did not affect the amount of ³²P transferred. These studies demonstrate that: (1) phosphorus is transferred among neighboring species in tallgrass prairie plant communities, (2) phosphorus may be transferred over significantly greater distances than reported in other grasslands, and (3) there is differential transfer among co-occurring species. Hypothesized mechanisms accounting for these patterns in tallgrass prairie include mycorrhizal hyphal interconnections and/or extensive and differential root and rhizosphere overlap among neighboring species.     

Nitrogen sink strength of ectomycorrhizal morphotypes of <Emphasis Type="Italic">Quercus douglasii</Emphasis>, <Emphasis Type="Italic">Q. garryana</Emphasis>, and <Emphasis Type="Italic">Q. agrifolia</Emphasis> seedlings grown in a northern California oak woodland

Little information is known on what the magnitude of nitrogen (N) processed by ectomycorrhizal (ECM) fungal species in the field. In a common garden experiment performed in a northern California oak woodland, we investigated transfer of nitrogen applied as ¹⁵NH₄ or ¹⁵NO₃ from leaves to ectomycorrhizal roots of three oak species, Quercus agrifolia, Q. douglasii, and Q. garryana. Oak seedlings formed five common ectomycorrhizal morphotypes on root tips. Mycorrhizal tips were more enriched in ¹⁵N than fine roots. N transfer was greater to the less common morphotypes than to the more common types. ¹⁵N transfer from leaves to roots was greater when , not , was supplied. ¹⁵N transfer to roots was greater in seedlings of Q. agrifolia than in Q. douglasii and Q. garryana. Differential N transfer to ectomycorrhizal root tips suggests that ectomycorrhizal morphotypes can influence flows of N from leaves to roots and that mycorrhizal diversity may influence the total N requirement of plants.     

Influence of arbuscular mycorrhizal fungi and simulated acid rain on the growth and coexistence of the grasses Calamagrostis villosa and Deschampsia flexuosa

Two grass species — Calamagrostis villosa (Chaix) J.F. Gmelin and Deschampsia flexuosa (L.) Trin. — are expanding in mountain Norway spruce (Picea abies L. Karst.) forests of Central Europe damaged by anthropogenic pollution constituted particularly of acid rain. This invasion of grasses may be caused by the higher irradiance reaching the forest floor after the pollution-induced tree defoliation. The relative abundance of the two grass species is changing during the process of forest decline. Our study investigated the effects of arbuscular mycorrhizal fungi (AMF) on the growth and coexistence of both species under simulated acid rain (SAR) and two levels of irradiance. Three microcosm experiments were conducted to investigate how both grasses are influenced by the AMF when grown separately or together interacting via extraradical mycelium (ERM). A positive growth response to inoculation with Glomus mosseae BEG 25 was found for both grass species when cultivated separately and the mycorrhizal dependence and the growth benefit for D. flexuosa was greater than for C. villosa. However, when both grass species were grown together in the rhizoboxes with separated root and hyphal compartments, the growth effect of the AMF was the opposite, i.e. C. villosa benefited more. The plants did not benefit from the AMF inoculation under the SAR treatment compared with dH2O treatment. The SAR also negatively influenced root length colonised by AMF, length of the ERM, alkaline phosphatase and NADH diaphorase activities of the ERM. The role of the ERM in transporting phosphorus between these grasses was verified by applying the radioisotope ³²P. There was a greater transport of isotopic ³²P between inoculated plants C. villosa and D. flexuosa grown in separated root compartments, as compared to non-inoculated plants. The amount of transported ³²P was low: a maximum of 3% of applied ³²P was detected in the shoots of receiver plants. Mechanical disturbance of the ERM significantly decreased the ³² P transport between plants. The ³²P transport between mycorrhizal plants was higher in the D. flexuosa to C. villosa direction than in the opposite one. Neither the SAR nor the low level of irradiance influenced the amount of transported ³²P. We discuss the role of ERM links between root systems in the coexistence of both grass species. This revised version was published online in June 2006 with corrections to the Cover Date.     

Influence of mycorrhizal associations on foliar δ15N values of legume and non-legume shrubs and trees in the fynbos of South Africa: Implications for estimating N2 fixation using the 15N natural abundance method

In this study, we examined the use of the ¹⁵N natural abundance method to quantify the percentage N derived from fixation of atmospheric N₂ in honeybush (Cyclopia spp.) shrubs and trees in the fynbos, South Africa. Non-fixing shrubs and trees of similar phenology to the Cyclopia species were chosen as reference plants. These reference plants were selected to cover a range of mycorrhizal associations (ericoid mycorrhizal, arbuscular mycorrhizal and non-mycorrhizal). Isotopic analysis revealed a wide range of foliar ¹⁵N values for the reference plants, including many very negative values. The marked differences in ¹⁵N values were defined by the mycorrhizal status of the reference plant species, with the ericoid and arbuscular mycorrhizal plants showing lower foliar ¹⁵N values relative to their non-mycorrhizal counterparts. In contrast, the ¹⁵N values of the N₂-fixing Cyclopia species were uniformly clustered around zero, from –0.11 to –1.43. These findings are consistent with the observation that mycorrhizal fungi discriminate against the heavier ¹⁵N isotope during transfer of N from the fungus to the host plant, leaving the latter depleted in ¹⁵N (i.e. with a more negative ¹⁵N value). However, a major assumption of the ¹⁵N natural abundance method for estimating N₂ fixation is that both legume and reference plant should have the same level of fractionation associated with N uptake. But, because mycorrhizal associations may strongly affect the level of fractionation during N uptake and transfer, the test legume should belong to the same mycorrhizal group as the chosen reference plant species. As shown in this study, if the mycorrhizal status of the legume and the reference plant differs, or cannot be assessed, then the ¹⁵N natural abundance technique cannot be used to quantitatively estimate N₂ fixation.     

Nitrogen nutrition,photosynthesis and carbon allocation in ectomycorrhizal pine

Summary Studies examined net photosynthesis (Pn) and dry matter production of mycorrhizal and nonmycorrhizalPinus taeda at 6 intervals over a 10-month period. Pn rates of mycorrhizal plants were consistently greater than nonmycorrhizal plants, and at 10 months were 2.1-fold greater. Partitioning of current photosynthate was examined by pulse-labelling with¹⁴CO₂ at each of the six time intervals. Mycorrhizal plants assimilated more¹⁴CO₂, allocated a greater percentage of assimilated¹⁴C to the root systems, and lost a greater percentage of¹⁴C by root respiration than did nonmycorrhizal plants. At 10 months, the quantity of¹⁴CO₂ respired by roots per unit root weight was 3.6-fold greater by mycorrhizal than nonmycorrhizal plants. Although the stimulation of photosynthesis and translocation of current photosynthate to the root system by mycorrhiza formation was consistent with the source-sink concept of sink demand, foliar N and P concentrations were also greater in mycorrhizal plants.Further studies examined Pn and dry matter production ofPinus contorta in response to various combinations of N fertilization (3, 62, 248 ppm), irradiance and mycorrhizal fungi inoculation. At 16 weeks of age, 6 weeks following inoculation with eitherPisolithus tinctorius orSuillus granulatus, Pn rates and biomass were significantly greater in mycorrhizal than nonmycorrhizal plants. Mycorrhizal plants had significantly greater foliar %P, but not %N, than did nonmycorrhizal plants. Fertilization with 62 ppm N resulted in greater mycorrhiza formation than either 3 or 248 ppm. Increased irradiance resulted in increased mycorrhiza formation.     

Partial mycoheterotrophy is common among chlorophyllous plants with Paris-type arbuscular mycorrhiza

Background and AimsAn arbuscular mycorrhiza is a mutualistic symbiosis with plants as carbon providers for fungi. However, achlorophyllous arbuscular mycorrhizal species are known to obtain carbon from fungi, i.e. they are mycoheterotrophic. These species all have the Paris type of arbuscular mycorrhiza. Recently, two chlorophyllous Paris-type species proved to be partially mycoheterotrophic. In this study, we explore the frequency of this condition and its association with Paris-type arbuscular mycorrhiza.MethodsWe searched for evidence of mycoheterotrophy in all currently published ¹³C, ²H and ¹⁵N stable isotope abundance patterns suited for calculations of enrichment factors, i.e. isotopic differences between neighbouring Paris- and Arum-type species. We found suitable data for 135 plant species classified into the two arbuscular mycorrhizal morphotypes.Key ResultsAbout half of the chlorophyllous Paris-type species tested were significantly enriched in ¹³C and often also enriched in ²H and ¹⁵N, compared with co-occurring Arum-type species. Based on a two-source linear mixing model, the carbon gain from the fungal source ranged between 7 and 93 % with ferns > horsetails > seed plants. The seed plants represented 13 families, many without a previous record of mycoheterotrophy. The ¹³C-enriched chlorophyllous Paris-type species were exclusively herbaceous perennials, with a majority of them thriving on shady forest ground.ConclusionsSignificant carbon acquisition from fungi appears quite common and widespread among Paris-type species, this arbuscular mycorrhizal morphotype probably being a pre-condition for developing varying degrees of mycoheterotrophy.