Invasion biology of Australian ectomycorrhizal fungi introduced with eucalypt plantations into the Iberian Peninsula

In the last two centuries, several species of Australian eucalypts (e.g. Eucalyptus camaldulensis and E.␣globulus) were introduced into the Iberian Peninsula for the production of paper pulp. The effects of the introduction of exotic root-symbitotic fungi together with the eucalypts have received little attention. During the past years, we have investigated the biology of ectomycorrhizal fungi in eucalypt plantations in the Iberian Peninsula. In the plantations studied, we found fruit bodies of several Australian ectomycorrhizal fungi and identified their ectomycorrhizas with DNA molecular markers. The most frequent species were Hydnangium carneum, Hymenogaster albus, Hysterangium inflatum, Labyrinthomyces donkii, Laccaria fraterna, Pisolithus albus, P. microcarpus, Rhulandiella berolinensis, Setchelliogaster rheophyllus, and Tricholoma eucalypticum. These fungi were likely brought from Australia together with the eucalypts, and they seem to have facilitated the establishment of eucalypt plantations and their naturalization. The dispersion of Australian fungal propagules may be facilitating the spread of eucalypts along watercourses in semiarid regions increasing the water lost. Because ectomycorrhizal fungi are obligate symbionts, their capacity to persist after eradication of eucalypt stands, and/or to extend beyond forest plantations, would rely on the possibility to find compatible native host trees, and to outcompete the native ectomycorrhizal fungi. Here we illustrate the case of the Australasian species Laccaria fraterna, which fruits in Mediterranean shrublands of ectomycorrhizal species of Cistus (rockroses). We need to know which other Australasian fungi extend to the native ecosystems, if we are to predict environmental␣risks associated with the introduction of Australasian ectomycorrhizal fungi into the Iberian Peninsula. This revised version was published online in July 2006 with corrections to the Cover Date.     

Assessment and monitoring of damage from insects in Australian eucalypt forests and commercial plantations

Abstract   This paper presents a review of recent developments in the assessment and monitoring of health in Australian eucalypt forests and plantations of pine and eucalypt species, with an emphasis on damage caused by herbivorous insects. The diverse range of interests and priorities amongst Australian stakeholders of native forests and plantations influences the scale, resolution and accuracy of results sought, and this in turn influences how the assessment data are collected, analysed and reported. The authors discuss sampling systems that include extensive ground-based surveys, permanent plots and airborne technologies being developed in Australia. In all cases, there is an appreciation that the assessment protocols should be objective, repeatable and cost effective. Significant progress has been made in the application of digital, remotely sensed imagery to detect and classify damaged forest canopies. The success of this approach depends, in part, on a sound understanding of the progression of symptoms at the leaf, tree crown and stand scale, especially those symptoms that influence spectral reflectance behaviour.     

Effects of P fertilisation and ectomycorrhizal fungal inoculation on early growth of eucalypt plantations in southern China

Eucalypt plantations in China have largely been established on soils that are low in phosphorus (P) and have few eucalypt-compatible ectomycorrhizal fungi. Effects of P application and ectomycorrhizal fungal inoculation on early tree growth in plantations of Eucalyptus urophylla Blake in Guangdong (Gaoyao) and E. globulus Labill. in Yunnan (Chuxiong) in southern China were investigated as part of a larger study. Application of superphosphate at establishment, in the presence of a basal fertiliser, increased early growth of E. urophylla and E. globulus. The optimum treatments for maximum stand volume at year 3 were 200 kg P ha^–1 which increased stand volume by 750% on the strongly acidic, P-deficient lateritic red oxisol at Gaoyao, and 40 kg P ha^–1 which increased stand volume by 55% on the mildly P-deficient red ultisol at Chuxiong, at 3 years. Superphosphate increased tree survival at Gaoyao as well as at Chuxiong. Nursery inoculation of eucalypt seedlings with ectomycorrhizal fungi significantly affected tree height and stand volume of the E. urophylla plantation, but the effect (positive or negative) was isolate-dependent and related to tree survival rate. A Laccaria isolate (CSIRO E4728) significantly increased stand volume by 27% at Gaoyao and a Scleroderma (MURU LH041) increased growth by 15% at Chuxiong at age 3 years. All isolates increased tree growth under P-limited soil conditions and only one isolate increased tree growth at marginal soil P. The results suggest that tree growth should be able to be optimised in plantations by the use of effective ectomycorrhizal fungi combined with a judicious fertilisation program at establishment.     

Consideration of the taxonomy and biodiversity of Australian ectomycorrhizal fungi

Mycorrhiza management in forestry must be predicated on an understanding of fungal biology and ecology. A fundamental building block of the biology and ecology of any organism is accurate identification and an understanding of its relationship to other organisms. The taxonomy of the larger fungi has been largely based on morphological classification of sexual structures but now Taxonomy routinely incorporates mating studies, and biochemical and molecular data. Taxonomy may not revolutionize theories on mycorrhiza but can clarify some of the inconsistencies due to misrepresentation or over-generalizations and inappropriate conclusions drawn from studies with inaccurately identified fungi. To illustrate this, we discuss and example where incorrect fungal names were repeatedly erroneously applied in morphological and physiological research reports on this fungus, e.g. Laccaria laccata. In this case subsequent taxonomic study revealed the reason for the conflicting research results reported for this fungus. We discuss the status of identifying the ectomycorrhizal fungi in various forest communities in Australia and the relationships of this process to assessing their use in forestry. Recent intensive efforts to collect, isolate and identify Australian ectomycorrhizal fungi have revealed an enormous and unique species diversity, e.g., for truffle-like fungi, over 2000 collections from the last five years alone have yielded 2 new families, 24 new genera, and about 184 new species. Nearly 95% of the described and undescribed fungi from Australia are novel, with some 22 genera and 3 families endemic. In most cases the current systematic knowledge of mycorrhizal fungi is inadequate to support clear framework for Australian taxa. This reflects the traditional Northern Hemisphere view of the world, the uniqueness of the Australian fungal flora, and how poorly it is known. For example, the genus Hymenogaster had been widely acknowledged as the most reduced member of the Cortinariaceae. However recent work on Hymenogaster species from the Southern Hemisphere has offered a number of alternative affinities to various species. We also discuss the role proper identification of the organisms involved plays in understanding the ecosystem. Emphasis should be placed on how species diversity equates with physiological and genetic diversity and how a sound taxonomic understanding of species and their systematic position is essential to properly manage them. Accurate taxonomic information will continue to be required as the basis for assessing the role of ectomycorrhizal fungi in sustained ecological development. Of particular significance is the role of ectomycorrhizal fungi in maintenance of plant diversity in natural ecosystems and those disturbed by management. In conclusion, we present some key research areas involving the use of taxonomy that need priority attention.     

Biological mobilization of potassium from clay minerals by ectomycorrhizal fungi and eucalypt seedling roots

Plant and Soil - Growth, proton and oxalate efflux, K&;nbsp;absorption and mineral depletion by one isolate of Cenococcum geophilum&;nbsp;Fr., by a putative isolate of Pisolithus microcarpus...     

Ecological role of hypogeous ectomycorrhizal fungi in Australian forests and woodlands

The Australian continent is characterised by a harsh climate and highly weathered, nutrient-poor soils. Trees and shrubs in these stressful environmental conditions typically form ectomycorrhizae with a variety of fungi, many of which form hypogeous (underground) fruit-bodies. The total number of hypogeous fungi Australia-wide is unknown, although recent systematic studies in the far south-eastern corner of the country suggest that they may number well over a thousand. Similar surveys elswhere are urgently required to clarify the situation. The precise ecological role of many hypogeous fungi remains to be determined, although most presumably facilitate nutrient and water uptake on behalf of their mycorrhizal partners. Others may also protect their plant host from root pathogens. One key function of hypogeous fungi is the role their fruit-bodies play as a food resource for a large range of terrestrial mammals. For a few animals, hypogeous fungi form the single most important dietary item year-round, whereas for others they may only be of seasonal or supplementary value. The extent to which fungi form part of the diet of any mammal species is reflected in the various levels of adaptation toward acquiring, then processing and digesting these cryptic and nutritionally challenging foodstuffs.     

Native and non-native mammals disperse exotic ectomycorrhizal fungi at long distances from pine plantations

We aimed to evaluate if exotic ectomycorrhizal fungi from exotic pine plantations disperse through non-native, but also native, mammals in a mountain ecosystem devoid of native ECM plants. Among four non-native and three native mammal species, feces of non-native wild boar (Sus scrofa) and brown hare (Lepus europaeus), and native pampa fox (Lycalopex gymnocercus) were selected to inoculate seedlings of Pinus elliottii. These feces came from two transects in an elevation gradient (1350–2250 m asl) and different distances from a pine plantation (100–6000 m). We show that feces of wild boar, brown hare (non-native), and pampa fox (native) were effective as inoculum for establishing ectomycorrhizal pine seedlings. Through molecular analyses, we determined that two species are mostly consumed and successfully form ectomycorrhizas with pine roots: Suillus granulatus and Rhizopogon pseudoroseolus. We provide novel evidence for the long-distance dispersal of exotic ectomycorrhizal fungi by non-native and native animal vectors.     

外生菌根真菌的共生互作和宿主选择机制研究进展

外生菌根真菌作为树木的共生伙伴,是森林生态系统重要组成部分,在森林天然更新、植物抗逆性形成、协助植物吸收限制性营养等方面扮演重要角色。真菌和植物跨界共生具有复杂的分子互作过程,在共生的不同阶段有不同的分子互作机制,其调控反馈网络还有许多未知。基因组与转录组研究技术和方法的进步,为一些新的信号分子、效应蛋白以及相关通路的发现提供了可能。真菌与宿主植物之间营养转移调控对共生的影响也逐渐受到关注,营养相关的转运蛋白对共生的建立和维持提供了物质基础。真菌的宿主选择机制是值得重点关注的领域,由于外生菌根真菌的多谱系起源和演化史中存在多次宿主转换事件,真菌演化出多样的应对机制用来区分相容性宿主、不相容性宿主和非宿主。通过对不同真菌与宿主植物的组学研究,宿主选择机制研究取得了一定进展。本文对近十年来国内外的研究报道进行梳理与总结,并对未来在该领域的探索方向做出展望。     

Options for managing chrysomelid leaf beetles in Australian eucalypt plantations: reducing the chemical footprint

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Phylogenetic determinants of potential host shifts in fungal pathogens

Understanding what determines the host range of pathogens and the potential for host shifts is of critical importance to controlling their introductions into new environments. The phylogeny of the hosts has been shown to be important: pathogens are more likely to be infectious on hosts closely related to their host‐of‐origin because of the similar host environments that is shared by descent. The importance of pathogen phylogenies for predicting host range has never been investigated, although a pathogen should also be able to exploit a new host that its close relative can infect. We performed cross‐inoculations using a plant–fungal association and showed that both host and pathogen phylogenies were significant predictors of host range, with at least partly independent effects. Furthermore, we showed that some pathogens were better at infecting novel hosts. Our results should have implications in the context of biological invasions and emergences of new diseases due to globalization.     

Zn pollution counteracts Cd toxicity in metal-tolerant ectomycorrhizal fungi and their host plant, Pinus sylvestris

Adaptive Zn and Cd tolerance have evolved in populations of the ectomycorrhizal fungus Suillus luteus. When exposed to high concentrations of both metals in vitro, a one-sided antagonism was apparent in the Zn- and Cd-tolerant isolates. Addition of high Zn concentrations restored growth of Cd-stressed isolates, but not vice versa. The antagonistic effect was not detected in a S. luteus isolate from non-contaminated land and in Paxillus involutus. The fungi were inoculated on pine seedlings and subsequently exposed to ecologically relevant Zn and Cd concentrations in single and mixed treatments. The applied doses severely reduced nutrient acquisition of non-mycorrhizal pines and pines inoculated with metal-sensitive S. luteus. Highest translocation of Zn and Cd to shoots occurred in the same plants. Seedlings inoculated with fungi collected from the polluted site reduced metal transfer to their host and maintained nutrient acquisition under high metal exposure. The isolate showing highest tolerance in vitro also offered best protection in symbiosis. The antagonistic effect of high Zn on Cd toxicity was confirmed in the plant experiment. The results indicate that a Zn- and Cd-polluted soil has selected ectomycorrhizal fungi that are able to survive and protect their phytobiont from nutrient starvation and excessive metal uptake.     

Fruiting of putative ectomycorrhizal fungi under blue gum (Eucalyptus globulus) plantations of different ages in Western Australia

 The species richness of putative ectomycorrhizal (EM) fungi fruiting in blue gum (Eucalyptus globulus Labill.) plantations in Western Australia was investigated in relation to plantation age. Eleven plantations, 1–8 years old, were selected for study and two native Eucalyptus forest sites in the same region were chosen for comparison. Sporocarps of 44 species of putative EM fungi were collected from the 13 sites. Of these, 30 species were found in blue gum plantations. The number of fungal species was highly positively correlated with plantation age and inversely correlated with soil pH. Young plantations (1–5 years) had 2–9 fungal species and were overwhelmingly dominated by species of Laccaria and Scleroderma. In older plantations (6–8 years), the relative abundance of sporocarps of each species within the fungal community decreased, accompanied by an increase in the number of fungal species (12–17 per site). A brief survey of the two native eucalypt forests in this region revealed a much higher number of fungal species than that observed in plantations. In plantations, species of Descolea, Laccaria, Pisolithus and Scleroderma typically fruited in young plantations. Species of epigeous fungi of the genera Boletus, Cortinarius, Hydnum, Inocybe, Lactarius, Paxillus, Russula and hypogeous fungi, including species of Descomyces, Hysterangium and Mesophellia, were found only in older plantations, or in native forests. Some of the fungi that fruit in young plantations are now being evaluated for use in commercial spore inoculation programs to increase the species diversity of EM fungi in exotic eucalypt plantations. Accepted: 8 October 1998     

Biomass and nutrient distribution in eucalypt plantations growing on rehabilitated bauxite mines

The quantities of above ground biomass and nutrients accumulated by two eucalypt forests growing on rehabilitated bauxite mines in south-western Western Australia were determined. The plantation at one site was 7.5 years old and had no understorey. The plantation at the second site was 3.5 years old and had been sown with seed of leguminous understorey species, resulting in a dense understorey. Similar amounts of biomass had accumulated at the two sites. However, the second site, in which the legume understorey accounted for 67% of the total above ground biomass, had about three times as much nitrogen in its biomass as the site without an understorey. There was also more sulphur in the biomass at this site. There were similar amounts of other nutrients at both sites. The plantation with an understorey is considered more likely to develop into a forest similar to the eucalypt forest prior to mining than the plantation without an understorey.     

外生菌根真菌对科尔沁沙地樟子松人工林衰退的影响

通过对科尔沁沙地东南缘樟子松人工林气温、地温及樟子松人工林根系分布的观测,从温度对外生菌根菌存活与生长影响的角度分析了沙地樟子松人工林衰退的现象,结果表明：由于沙地表层土壤（0～5cm）高温环境,外生菌根菌无法存活;在沙地20～40cm土层内,温度条件有利于外生菌根菌的存活、生长,而13～42年生沙地樟子松人工林树木平均78％的根系分布于该层,说明外生菌根菌的存在阻止或延缓了樟子松人工林的生长衰退,不是导致沙地樟子松人工林枯叶、枯梢、枯枝、死亡的原因．表层土壤外生菌根菌的缺乏使萌发的幼苗不能形成有效根系是导致沙地樟子松人工林不能天然更新的主要原因之一．     

Genetics of ectomycorrhizal fungi: progress and prospects

The variability within and among ectomycorrhizal species provides a substantial genetic resource and the potential to increase forest productivity and environmental sustainability. Two parallel and interacting approaches, classical and molecular genetics, are being developed to acquire the genetic information underpinning selection of improved ectomycorrhizal strains. Determining the genetic traits of the fungi which contribute to symbiosis and plant function are being followed using natural variability combined with classical and molecular genetic manipulations. Classical and molecular manipulations for breeding rely on key information including sexual and parasexual reproduction, postmeiotic nuclear behaviour, mating-types and vegetative incompatibility mechanisms. Progress in the manipulation of genomes of ectomycorrhizal fungi will depend on efficient methods for gene cloning and DNA transformation. Gene transfer into fungal cells have been shown to be successful and include treatment of protoplasts and intact mycelium with naked DNA in the presence of polyvalent cations, electroporation, and microbombardment. The merits and limitations of these methods are discussed. Using this technology the expression of foreign DNA, the functional analysis of fungal DNA sequences, as well as molecular exploitation for commercial purposes can be carried out. This review concentrates on these aspects of fungal molecular biology and discusses the applications of the experimental systems that are currently available to ectomycorrhizal fungi. As it is essential to be able to define the traits which a breeder is seeking to improve, availability of genetically defined strains that are isogenic for a character or differ only in one character and a thorough knowledge of the biochemistry of the symbiosis will be necessary before any genetic manipulation be carried out. Genetic variability of ectomycorrhizal strains has been assessed by DNA fingerprinting. This approach allows the evaluation of DNA variability and the exchange of genetic information in natural populations, the identification of species and isolates by DNA polymorphisms, and tracking the environmental fate of the introduced fungi to determine their survival, growth, and dissemination within the soil.     

Extramatrical structures of hydrophobic and hydrophilic ectomycorrhizal fungi

The extramatrical mycelia of Suillus bovinus, Rhizopogon luteolus and R. vinicolor, all examples of hydrophobic (ho), mat-forming mycorrhizal fungi, were examined while associated with their hosts in the unsterilized rhizoscope, and efforts were made to produce and examine similar structures in vitro. Comparisons were made with four hydrophilic (hi) mycorrhizal fungi, Thelephora terrestris, Cenococcum geophilum, Laccaria laccata and Hebeloma crustuliniforme. The ho fungi formed linear structures (coarse, rhizomorph-like cords, with vessels in the center) and fans, both in the rhizoscope and in vitro. The same was seen in mycorrhizal mycelia in forest soils. These cords did not themselves give rise to the fans peripherally, and were not proper rhizomorphs, but were created continuously from single exploring air hyphae in the preexisting fan. Thus the ho exploring hyphae aggregated into strands, which grew in thickness only when no suitable, exploitable substrate was found. The assembly of hyphae creating ho cords was seen in the air as well as on inert hydrophilic (glass) or hydrophobic (plastic) surfaces, but never in water. It is hypothesized that the ho cell wall surface glues hyphae together while cords are formed. Water disturbed strands and mantles already formed. The ho exploring hyphae could also create ho mycelial patches (as in a mat) at the water-air interface of a number of substrates. The periphery of these patches seemed to be composed of shorter exploiting hyphae penetrating different water-soaked substrates. Exploring, aerial hyphal tips of the ho fungi were shown to excrete water droplets from openings in the ho cell wall surface, both in vitro and in the rhizoscope. In the rhizoscope, droplet excretion was apparently directly governed by photosynthesis in the shoot of the seedling. It is proposed that the drop exudation represents a kidney-like function of the extramatrical hyphae and a bridge to drier soil particles to initiate nutrient uptake by the hyphae. The ecological function of the different extramatrical structures of ho fungi are discussed. The ho cords or hyphae may translocate water only in the vessels or symplastically and not in the cell walls. The ho property may be essential among the S-selected (stress-tolerant) factors in these forest fungi. The transfer from water-repelling exploring structures into more hi exploiting structures in water contact with surrounding soil debris is, therefore, of great importance. The hi fungi did not form rhizomorph-like strands, in most cases, but an extending hyphal mycelium, representing foraging, exploring and exploiting structures at the same time. In the field, short strands may be found. On the hi fungi droplets were also produced but readily fused into a water sheath around the hypha. The hyphae thus tended to wick water via the cell wall.