Cadmium immobilization in the rhizosphere of arbuscular mycorrhizal plants by the fungal extraradical mycelium

The aim of the study was to assess how the extraradical mycelium (ERM) of arbuscular mycorrhizal (AM) fungi contributes to Cd immobilization in the rhizosphere. Substrates prepared by cultivation of AM and non-mycorrhizal tobacco (Nicotiana tabacum L.) in quartz sand in two experiments were amended with Cd in a range of concentrations and Cd immobilization was assessed as Cd toxicity using root growth tests. Split-root plants, inoculated at one part of the root system, and hyphal compartments colonized by ERM only were used to separate the effects of ERM from plant-mediated effects of mycorrhiza and from the effects of roots. AM decreased Cd toxicity in the substrates obtained by 12 weeks of cultivation (Experiment 1), while the effect was less clear after 8 weeks (Experiment 2). No indication was found for an involvement of plant-mediated effects; in contrast, the effect of ERM could be clearly demonstrated. Lower Cd toxicity in the substrates colonized by ERM was related to ERM-induced alkalinization, but not directly to ERM density. It is concluded that the ERM of AM fungi may enhance Cd immobilisation in soil not only due to its high Cd sorption capacity but also by its activity.     

Associations of dominant plant species with arbuscular mycorrhizal fungi during vegetation development on coal mine spoil banks

Among plants colonizing mine spoil banks in Northern Bohemia the first colonizers, mainly ruderal annuals fromChenopodiaceae andBrassicaceae were found not to be associated with arbuscular mycorrhizal fungi (AMF). These species cultivated in pots with soil from four sites in different succession stages of the spoil bank did not respond to the presence of native or non-native AMF. All grass species studied (Elytrigia repens, Calamagrostis epigejos andArrhenatherum elatius) were found moderately colonized in the field and they seemed to be facultative mycotrophs, because their response to mycorrhiza in the experiment was negligible.Carduus acanthoides was found to be highly colonized in the field, however, it did not show growth response to AMF in the pot experiment. The AMF native in four sites on the spoil banks showed high infectivity but low effectiveness in association with colonizing plants compared to the non-native isolateG. fistulosum BEG23. In general, dependence on AMF in the cultivation experiment was rather low, regardless of the fact that plants were found to be associated with AMF either in the field or in pots. Occurrence and effectiveness of mycorrhizal associations might relate primarily to the mycotrophic status of each plant species rather than to the age of the spoil bank sites studied.     

How do Mycorrhizas Affect C and N Relationships in Flooded Aster tripolium Plants?

The mycorrhizal associations established between plants and fungi have multiple effects on plant growth, directly affecting stress tolerance. This work aimed to explore arbuscular mycorrhizal (AM) effects on carbon and nitrogen relationships of Aster tripolium L. and consequently on its flooding tolerance. Mycorrhizal and non-mycorrhizal juvenile plants were submitted to non-flooding and tidal flooding conditions for 56 d. Tidal flooding reduced biomass, but the presence of mycorrhiza had an ameliorating effect. The AM symbioses seem to have, like flooding, a stressful effect on A. tripolium at an early stage of plant development. However, once the plant was established, an improvement of growth performance of plants with mycorrhiza under flooding conditions was observed. The better tolerance of AM plants to flooding was mediated through an improvement of the osmotic adjustment of the plant tissues (higher concentrations of soluble sugars and proline) and through the increment of nitrogen acquisition in tidal-flooded plants.     

Does the sequence of plant dominants affect mycorrhiza development in simulated succession on spoil banks?

The aim of this field study was to examine how the development of arbuscular mycorrhizal fungi (AMF) on coal mine spoil banks is affected by the presence of plants with different mycorrhizal status. A 3-year trial was conducted on the freshly created spoil bank Vršany, North-Bohemian coal basin, the Czech Republic. Three plant species – non-mycotrophic annual Atriplex sagittata, highly mycotrophic annual Tripleurospermum inodorum (both dominants of early stages of succession) and facultatively mycotrophic Arrhenatherum elatius (a perennial grass species of the later stage of succession) – were planted on 1 m² plots over 3 years in different sequences that simulated the progress of succession on spoil banks. The development of AMF populations was monitored by evaluation of mycorrhizal colonization of plant roots and by measurement of the mycorrhizal inoculation potential (MIP) of soil. These two parameters were compared between plots inoculated with the mixture of three AMF isolates – Glomus mosseae BEG95, G. claroideum BEG96 and G. intraradices BEG140 – (“inoculated plots”) and plots exposed only to natural dispersal of AMF propagules (“uninoculated plots”). Highly colonized roots of plants together with a high MIP of soil in uninoculated plots were already found at the end of the first season, indicating rapid natural dispersal of AMF propagules. Root colonization of facultatively mycotrophic and non-mycotrophic plants in later years was affected by the mycorrhizal status of the previous plant species. The MIP of soil continuously increased throughout the experiment; in uninoculated plots, the MIP was temporarily decreased if plant species of higher mycotrophy were replaced by species of lower mycotrophy. The results lead to the conclusion that AMF colonize freshly formed sites very quickly and reproduce or accumulate in the soil, which leads to increasing MIP values. However, this infective potential can be decreased if non-mycotrophic plants predominate on the site.     

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.     

葡萄糖、根浸出液对丛枝菌根真菌吸收不同外源氮产生精氨酸的影响

以大葱(Allium fistulosum)为宿主植物, 接种丛枝菌根(Arbuscular mycorrhizal, AM)真菌Glomus intraradices, 采用三室隔离盆栽培养系统, 在菌丝室施加浓度为4 mmol/L的不同形态外源氮、1%葡萄糖及根浸出液, 通过测定根外菌丝(Extraradical mycelium, ERM)和菌根中精氨酸的含量, 探究葡萄糖、根浸出液对AM真菌吸收不同形式外源氮产生精氨酸的影响。结果表明, 不同外源氮对ERM中精氨酸含量的影响为尿素>Gln>NH₄NO₃>Arg/Gly>NH₄Cl>KNO₃, 对菌根中精氨酸含量的影响为Arg>Gln>尿素>NH₄NO₃>Gly>NH₄Cl>KNO₃; 施加葡萄糖和根浸出液在不同程度上提高ERM干重和菌丝室孢子数量, 但使ERM和菌根中的精氨酸含量降低。说明AM真菌吸收同化不同外源氮产生精氨酸的能力不同, 葡萄糖和根浸出液降低AM真菌吸收同化外源氮产精氨酸的能力。     

Transport of nitrogen and zinc to rhodes grass by arbuscular mycorrhiza and roots as affected by different nitrogen sources (NH<Subscript>4</Subscript><Superscript>+</Superscript>-N and NO<Subscript>3</Subscript><Superscript>−</Superscript>-N)

We investigated the effect of mineral nitrogen forms on transfer of nitrogen (N) and zinc (Zn) from attached compartments to rhodes grass (Chloris gayana) colonised with arbuscular mycorrhizal fungi (AMF). After being pre-cultivated in substrates with adequate nutrient supply and either AMF inoculated (+AM) or left non-inoculated (?AM), rhodes grass was positioned adjacent to an outer compartment holding a similar substrate but applied with labelled nitrogen (¹⁵N) either as ammonium (NH₄ ⁺) or nitrate (NO₃ ^?), and a high supply of Zn (150 mg kg^?1 DS). Plant roots together with fungal mycelium were either allowed to explore the outer compartment (with root access) or only mycorrhizal hyphae were allowed (without root access). Within each access treatment, biomasses of rhodes grass were not significantly affected by AMF inoculation or N form. AMF contribution to plant ¹⁵N uptake was about double in NH₄ ⁺ compared with NO₃ ^?-supplied treatments while the mycorrhizal influence on plant Zn uptake was insignificant. Without root access, the shoot ¹⁵N/Zn concentration ratio was up to ten-fold higher in +AM than –AM treatments and this ratio increase was clearly more pronounced in NH₄ ⁺ than NO₃ ^?-supplied treatments. In conclusion, rhodes grass in symbiosis with the tested AMF acquired more N when supplied with ammonium. Moreover, there is clear indication that although the AMF have transported both nutrients (N and Zn), N was preferentially transferred as compared to Zn. We confirmed that, while rhodes grass is not able to prevent excessive Zn uptake via roots under conditions of high Zn, mycorrhiza is able to avoid excessive Zn supply to the host plant when the fungus alone has access to contaminated patches.     

Growth and viability of mycorrhizal extraradical mycelia associated with three temperate orchid species

Growth and enzymatic activities of extraradical mycelia (ERM) of native mycorrhizal symbionts associated with three orchid species, Dactylorhiza fuchsii, D. majalis and Platanthera bifolia, were studied. ERM extracted from the mycorrhizosphere of these species showed features typical for fungi that form orchid mycorrhiza. In the first pot experiment, three different treatments were applied on tubers of D. fuchsii transplanted from a natural site: control (no specific treatment), reinoculated (surface-sterilized tubers reinoculated with mycorrhizal fungi-colonised roots), and benomyl (nonsterilized tubers treated with fungicide). However, no significant differences in ERM growth and intensity of root mycorrhizal colonisation at harvest were observed among these treatments. ERM associated with reinoculated D. fuchsii plants showed significantly higher alkaline phosphatase (ALP) enzymatic activity at week 36 than at week 24, but no differences were observed for NADH diaphorase activity. Benomyl application significantly reduced ALP activity in comparison with reinoculated plants at week 36. In the second experiment, plants of all three species were either untreated (control), or repeatedly treated with benomyl. Similarly to the results of the first experiment, benomyl application did not reduce the ERM growth of mycorrhizal symbionts associated with D. majalis and D. fuchsii. The low ERM growth associated with benomyl-treated P. bifolia was probably caused by poor root system development in this treatment. Significantly higher mycorrhizal colonisation was found for D. fuchsii compared to P. bifolia in control treatments at the end of cultivation. The ERM of native symbionts of the three orchid species studied seemed to have a different growth pattern over time and responded differently to fungicide application.     

Mycorrhizal Networks Interacting with Litter Improves Nutrients and Growth for One Plant through the Vary of N/P Ratio under Karst Soil

Arbuscular mycorrhizae (AM) fungi affect nutrient uptake for host plants, while it is unclear how AM fungi interacting with soil litter affect plant growth and nutrient utilization through mycorrhizal networks in karst soil of deficient nutrients beyond the rhizosphere. An experiment was conducted in a microcosm composed of a planting compartment for Cinnamomum camphora seedlings with or without Glomus mosseae fungus (M⁺ vs. M⁻ ) and an adjacent litter compartment containing or not containing additional litter material of Arthraxon hispidus (L⁺ vs. L⁻ ), where the compartments are connected either by nylon mesh of 20 μm or 0.45 μm which either allow available mycorrhizal networks within the litter compartment or prevent mycelium entering into the litter compartment (N⁺ vs. N⁻ ). Plant biomass and nutrients were measured. The results showed that the addition of litter changed the symbiotic process in mycorrhizal colonization, spore, and hyphal density, which when in association with the host plant then affected the biomass, and accumulations of N (nitrogen) and P (phosphorus) in the individual plant as well as root, stem, and leaf respectively. AM fungi increased N and P accumulations and N/P ratio in individual plants and plant tissues. A decrease of the N/P ratio of the individual plant was observed when AM fungus interacted significantly with litter through mycorrhizal networks in the litter compartment. The results indicate that the C. camphora seedlings benefited from litter in nutrient utilization of N and P through the vary of N/P ratio when accessing mycorrhizal networks. These findings suggest that mycorrhizal networks interacting with litter improve growth and nutrients of N and P for plants through the vary of N/P ratio in order to alleviate nutrient limitation under karst soil.     

Effect of arbuscular mycorrhiza on inter- and intraspecific competition of two grassland species

We were interested in the role of arbuscular mycorrhiza (AM) in the competition between plants of different sizes. A pot experiment of factorial design was established, in which AM root colonization and competition were used as treatments. Five-week-old Prunella vulgaris seedlings were chosen as target plants (i.e. plants whose response to competition was studied) and the following (13 replicates of each) were used as neighbours: (1) a large, 10-week-old P. vulgaris, (2) two P. vulgaris seedlings, and (3) a large, 10-week-old Fragaria vesca. In the experiment where small neighbours were grown together with small target plants, competition did not reduce target plant weight significantly, compared to the other two treatments. The competitive effects of large neighbours were significant, regardless of species (both older neighbours reduced the weights of target plants similarly), but there was a clear difference between intra- and interspecific competition when plants were mycorrhizal. In intraspecific competition with a large neighbour, the target plant shoot weight was reduced 24% when inoculated with AM. Thus, AM amplified rather than balanced intraspecific competition. In interspecific competition with old F. vesca, the shoot weights of target plants were 22% greater when inoculated with AM than when non-mycorrhizal. The results showed that, for given soil condition, AM might increase species diversity by increasing competitive intraspecific suppression and decreasing the interspecific suppression of small plants by larger neighbours.     

Systemic inhibition of arbuscular mycorrhiza development by root exudates of cucumber plants colonized by<Emphasis Type="Italic"> Glomus mosseae</Emphasis>

The arbuscular mycorrhizal (AM) non-host plants mustard, sugar beet, lupin and the AM host plant cucumber were used as test plants. Cucumber plants were grown either in the absence of the AM fungus (AMF) Glomus mosseae or in a split-root system, with one side mycorrhizal and one side non-mycorrhizal. Root exudates of the AM non-host plants, the non-mycorrhizal cucumber plants and the mycorrhizal and the non-mycorrhizal side of the split-root system of mycorrhizal cucumber plants were collected and applied to cucumber plants inoculated with the AMF. Root exudates of non-mycorrhizal cucumber plants showed a significant stimulatory effect on root colonization, whereas root exudates from the mycorrhizal and the non-mycorrhizal sides of a split-root system of a mycorrhizal cucumber plant did not show this stimulatory effect and were even slightly inhibitory. Root exudates of the two AM non-host plants mustard and sugar beet significantly reduced root colonization in cucumber plants, whereas no such effect was observed when root exudates of the AM non-host plant lupin were applied.     

丛枝菌根对喜树幼苗生长和氮、磷吸收的影响

喜树(Camptotheca acuminata)是我国特有的多年生亚热带落叶阔叶树种,因其次生代谢产物喜树碱具有良好的抗肿瘤活性而受到人们的广泛关注。该文通过温室盆栽接种试验,观察了2属6种丛枝菌根真菌即蜜色无梗囊霉(Acaulospora mellea)、光壁无梗囊霉(A. laevis)、木薯球囊霉(Glomus manihot)、地表球囊霉(G. versiforme)、幼套球囊霉(G. etunicatum)和透光球囊霉(G. diaphanum)对喜树幼苗生长和氮、磷养分吸收的影响。结果表明,丛枝菌根的形成对喜树幼苗的生长以及氮、磷营养的吸收均有影响。从生物量看,除幼套球囊霉和光壁无梗囊霉侵染形成的丛枝菌根喜树幼苗与无菌根幼苗(CK)差异不显著外,其余菌根幼苗的生物量均明显大于无菌根幼苗,透光球囊霉和蜜色无梗囊霉菌根幼苗尤为突出,分别达到无菌根幼苗的1.9和1.4倍。丛枝菌根的形成似乎不利于喜树幼苗的氮素营养吸收,并且主要体现在叶片的氮含量上。相反,丛枝菌根形成总体上促进喜树幼苗对磷素营养的吸收,并且主要体现在根的磷含量上。与无菌根幼苗比,所有菌根幼苗根的氮、磷分配比例增加,而叶片的氮、磷分配比例减少。     

Interactions between plants: the role of mycorrhizae

Summary We present and discuss evidence, mostly from our own research, on some possible roles of mycorrhizae in interactions between plants. Experiments investigating whether seedlings become more rapidly infected with mycorrhiza if they are near large, already infected plants have shown that contact between the seedling's roots and established mycelium sometimes speeds up infection but on other occasions it does not. The reason for the discrepancies is not clear. Mycorrhiza can substantially alter the balance between competing plant species in a way that would not be predicted from their response when growing separately. An experiment involving large and small plants of the same species growing together showed little effect of mycorrhiza on the balance between plants of different sizes. The rate of transfer of ³²P between plants of Lolium perenne or Plantago lanceolata was so slow, even when they were mycorrhizal, that phosphorus transfer between living plants seems unlikely to be of major ecological importance. However, nitrogen was found to be transferred much more freely than phosphorus between P. lanceolata plants. Situations are discussed in which there could be a source-sink relationship between plants causing net flow of carbon or mineral nutrients from one to the other. If nutrients pass from dying roots to living plants via mycorrhizal links, this could result in preferential nutrient cycling between species that share the same type of mycorrhiza. Some evidence is presented that this does happen.     

Mycorrhizal mediation of plant N acquisition and residue decomposition: Impact of mineral N inputs

Mycorrhizas are ubiquitous plant–fungus mutualists in terrestrial ecosystems and play important roles in plant resource capture and nutrient cycling. Sporadic evidence suggests that anthropogenic nitrogen (N) input may impact the development and the functioning of arbuscular mycorrhizal (AM) fungi, potentially altering host plant growth and soil carbon (C) dynamics. In this study, we examined how mineral N inputs affected mycorrhizal mediation of plant N acquisition and residue decomposition in a microcosm system. Each microcosm unit was separated into HOST and TEST compartments by a replaceable mesh screen that either prevented or allowed AM fungal hyphae but not plant roots to grow into the TEST compartments. Wild oat (Avena fatua L.) was planted in the HOST compartments that had been inoculated with either a single species of AM fungus, Glomus etunicatum, or a mixture of AM fungi including G. etunicatum. Mycorrhizal contributions to plant N acquisition and residue decomposition were directly assessed by introducing a mineral ¹⁵N tracer and ¹³C‐rich residues of a C₄ plant to the TEST compartments. Results from ¹⁵N tracer measurements showed that AM fungal hyphae directly transported N from the TEST soil to the host plant. Compared with the control with no penetration of AM fungal hyphae, AM hyphal penetration led to a 125% increase in biomass ¹⁵N of host plants and a 20% reduction in extractable inorganic N in the TEST soil. Mineral N inputs to the HOST compartments (equivalent to 5.0 g N m^?2 yr^?1) increased oat biomass and total root length colonized by mycorrhizal fungi by 189% and 285%, respectively, as compared with the no‐N control. Mineral N inputs to the HOST plants also reduced extractable inorganic N and particulate residue C proportion by 58% and 12%, respectively, in the corresponding TEST soils as compared to the no‐N control, by stimulating AM fungal growth and activities. The species mixture of mycorrhizal fungi was more effective in facilitating N transport and residue decomposition than the single AM species. These findings indicate that low‐level mineral N inputs may significantly enhance nutrient cycling and plant resource capture in terrestrial ecosystems via stimulation of root growth, mycorrhizal functioning, and residue decomposition. The long‐term effects of these observed alterations on soil C dynamics remain to be investigated.     

Use of Arbuscular Mycorrhiza and Organic Amendments to Enhance Growth of Macaranga peltata (Roxb.) Müll. Arg. in Iron Ore Mine Wastelands

Macaranga peltata (Roxb.) Mull. Arg. is a disturbance tolerant plant species with potential in mine wasteland reclamation. Our study aims at studying the phyto-extraction potential of M. peltata and determining plant-soil interaction factors effecting plant growth in iron ore mine spoils. Plants were grown in pure mine spoil and spoil amended with Farm Yard Manure (FYM) and Vermicompost (VC) along with arbuscular mycorrhizal (AM) species Rhizophagus irregularis. Pure and amended mine spoils were evaluated for nutrient status. Plant growth parameters and foliar nutrient contents were determined at the end of one year. FYM amendment in spoil significantly increased plant biomass compared to pure mine spoil and VC amended spoil. Foliar Fe accumulation was recorded highest (594.67μg/g) in pure spoil with no mortality but considerably affecting plant growth, thus proving to exhibit phyto-extraction potential. FYM and VC amendments reduced AM colonization (30.4% and 37% resp.) and plants showed a negative mycorrhizal dependency (–30.35 and –39.83 resp.). Soil pH and P levels and, foliar Fe accumulation are major factors determining plant growth in spoil. FYM amendment was found to be superior to VC as a spoil amendment for hastening plant growth and establishment in iron ore mine spoil     

Mycorrhiza-plant colonization patterns on a subalpine glacier forefront as a model system of primary succession

Lyman glacier in the North Cascades Mountains of Washington has a subalpine forefront characterized by a well-developed terminal moraine, inconspicuous successional moraines, fluting, and outwash. These deposits were depleted of symbiotic fungi when first exposed but colonized by them over time after exposure. Four major groups of plant species in this system are (1) mycorrhiza-independent or facultative mycotrophic, (2) dependent on arbuscular mycorrhizae (AM) (3) dependent on ericoid mycorrhiza (ERM) or ectomycorrhizae (EM), and (4) colonized by dark-septate (DS) endophytes. We hypothesized that availability of mycorrhizal propagules was related to the success of mycorrhiza-dependent plants in colonizing new substrates in naturally evolved ecosystems. To test this hypothesis roots samples of 66 plant species were examined for mycorrhizal colonization. The plants were sampled from communities at increasing distances from the glacier terminus to compare the newest communities with successively older ones. Long established, secondary successional dry meadow communities adjacent to the glacier forefront, and nearby high alpine communities were sampled for comparison. DS were common on most plant species on the forefront. Nonmycorrhizal plants predominated in the earlier successional sites, whereas the proportion of mycorrhizal plants generally increased with age of community. AM were present, mostly at low levels, and nearly absent in two sites of the forefront. ERM were present in all species of Ericaceae sampled, and EM in all species of Pinaceae and Salicaceae. Roots of plants in the long established meadow and heath communities adjacent to the forefront and the high alpine community all had one or another of the colonization types, with DS and AM predominating.     

Effect of<Emphasis Type="Italic">Glomus intraradices</Emphasis> isolated from Pb-contaminated soil on Pb uptake by<Emphasis Type="Italic">Agrostis capillaris</Emphasis> is changed by its cultivation in a metal-free substrate

Development and heavy metal tolerance of two cultivation lineages of the indigenous isolate of arbuscular mycorrhizal fungus (AMF)Glomus intraradices PH5 were compared in a pot experiment in soil from lead (Pb) smelter waste deposits. One lineage was sub-cultured in original Pb-contaminated soil; the second one was maintained for 13 months in an inert substrate (river sand) without Pb stress. The contribution of these cultivation lineages to the Pb uptake and accumulation by the host plantAgrostis capillaris was investigated. The experiment was conducted in a compartmented system where the lateral compartments withAgrostis seedlings were separated from the central pot containing 4-week olderAgrostis plants by a nylon mesh for allowing out-growing of extraradical mycelium (ERM) from the pot. No differences in mycorrhizal colonization, ERM length and viability were observed between the two lineages ofG. intraradices PH5 in the soil of the isolate origin. However, the ability to support plant growth and Pb uptake differed between the lineages and also between the plants in the central pots and the lateral compartments. The growth of the plants in the central pots was positively affected by AMF inoculation. The plants inoculated with the lineage maintained in original soil showed larger shoot biomass and higher shoot P content as compared to the other inoculation treatments. The shoot Pb concentration of these plants was lower when compared to the plants inoculated with the lineage sub-cultured in the inert substrate. However the concentration did not differ from non-mycorrhizal control or from the reference isolateG. intraradices BEG75 from non-contaminated soil. Also shoot Pb contents were similar for all inoculation treatments. The development ofG. intraradices BEG75 in the contaminated soil was very poor; this isolate was not able to initiate colonization of seedlings in lateral compartments. In lateral compartments, growth of seedlings in contaminated soil was inhibited by theG. intraradices PH5 lineage maintained in the inert substrate. Pb translocation from the seedling roots to shoots was increased for plants inoculated with either lineage as compared to the non-mycorrhizal control; however, the increase for the lineage cultivated in the inert substrate was significantly higher in comparison with that maintained in the original soil. After 13 months of cultivation in a metal free substrate, theG. intraradices isolate from Pb contaminated soil did not lose its tolerance to Pb as regards colonization of plant roots and growth of ERM in the soil of its origin. However, its ability to support plant growth and to prevent Pb translocation from the roots to the shoots was decreased.     

Functional aspects of root architecture and mycorrhizal inoculation with respect to nutrient uptake capacity

The aim of this research was to investigate the effect of arbuscular mycorrhizal (AM) colonisation on root morphology and nitrogen uptake capacity of carob ( Ceratonia siliqua L.) under high and low nutrient conditions. The experimental design was a factorial arrangement of presence/absence of mycorrhizal fungus inoculation ( Glomus intraradices) and high/low nutrient status. Percent AM colonisation, nitrate and ammonium uptake capacity, and nitrogen and phosphorus contents were determined in 3-month-old seedlings. Grayscale and colour images were used to study root morphology and topology, and to assess the relation between root pigmentation and physiological activities. AM colonisation lead to a higher allocation of biomass to white and yellow parts of the root. Inorganic nitrogen uptake capacity per unit root length and nitrogen content were greatest in AM colonised plants grown under low nutrient conditions. A better match was found between plant nitrogen content and biomass accumulation, than between plant phosphorus content and biomass accumulation. It is suggested that the increase in nutrient uptake capacity of AM colonised roots is dependent both on changes in root morphology and physiological uptake potential. This study contributes to an understanding of the role of AM fungi and root morphology in plant nutrient uptake and shows that AM colonisation improves the nitrogen nutrition of plants, mainly when growing at low levels of nutrients.     

Root Anatomy and Mycotrophy (AM) of the Achlorophyllous Voyria truncata (Standley) Standley & Steyermark (Gentianaceae)

Roots of Voyria truncata retain the primary root structure even though they can grow as thick as 2 mm in diameter. These root diameters are due to a retained capability for cell division in the cortex parenchyma. This is explained as a vital adaptation to its life form. Based on the extraradical mycelium, the mode of penetration, the structurally incompatible intraradical phase, the presence of intercellular vesicles in the root cortex, and the occurrence of immediate hyphal bridges from arbuscular mycorrhizal roots of neighbouring plants, the mycorrhiza of V. truncata is described as an arbuscular mycorrhiza (AM), although the characteristic arbuscles are missing. Special features of the AM in V. truncata are interpreted as an improved efficiency in taking advantage of the mycorrhiza. Root connections with roots of neighbouring plants are common and preferred locations for fungal infections. An evolutionary tendency towards parasitism of higher plants is discussed.