How belowground interactions contribute to the coexistence of mycorrhizal and non-mycorrhizal species in severely phosphorus-impoverished hyperdiverse ecosystems

Background

Mycorrhizal strategies are very effective in enhancing plant acquisition of poorly-mobile nutrients, particularly phosphorus (P) from infertile soil. However, on very old and severely P-impoverished soils, a carboxylate-releasing and P-mobilising cluster-root strategy is more effective at acquiring this growth-limiting resource. Carboxylates are released during a period of only a few days from ephemeral cluster roots. Despite the cluster-root strategy being superior for P acquisition in such environments, these species coexist with a wide range of mycorrhizal species, raising questions about the mechanisms contributing to their coexistence.

Scope

We surmise that the coexistence of mycorrhizal and non-mycorrhizal strategies is primarily accounted for by a combination of belowground mechanisms, namely (i) facilitation of P acquisition by mycorrhizal plants from neighbouring cluster-rooted plants, and (ii) interactions between roots, pathogens and mycorrhizal fungi, which enhance the plants’ defence against pathogens. Facilitation of nutrient acquisition by cluster-rooted plants involves carboxylate exudation, making more P available for both themselves and their mycorrhizal neighbours. Belowground nutrient exchanges between carboxylate-exuding plants and mycorrhizal N₂-fixing plants appear likely, but require further experimental testing to determine their nutritional and ecological relevance. Anatomical studies of roots of cluster-rooted Proteaceae species show that they do not form a complete suberised exodermis.

Conclusions

The absence of an exodermis may well be important to rapidly release carboxylates, but likely lowers root structural defences against pathogens, particularly oomycetes. Conversely, roots of mycorrhizal plants may not be as effective at acquiring P when P availability is very low, but they are better defended against pathogens, and this superior defence likely involves mycorrhizal fungi. Taken together, we are beginning to understand how an exceptionally large number of plant species and P-acquisition strategies coexist on the most severely P-impoverished soils.

     

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.     

Phosphorus-mobilization ecosystem engineering: the roles of cluster roots and carboxylate exudation in young P-limited ecosystems

Background

Carboxylate-releasing cluster roots of Proteaceae play a key role in acquiring phosphorus (P) from ancient nutrient-impoverished soils in Australia. However, cluster roots are also found in Proteaceae on young, P-rich soils in Chile where they allow P acquisition from soils that strongly sorb P.

Scope

Unlike Proteaceae in Australia that tend to proficiently remobilize P from senescent leaves, Chilean Proteaceae produce leaf litter rich in P. Consequently, they may act as ecosystem engineers, providing P for plants without specialized roots to access sorbed P. We propose a similar ecosystem-engineering role for species that release large amounts of carboxylates in other relatively young, strongly P-sorbing substrates, e.g. young acidic volcanic deposits and calcareous dunes. Many of these species also fix atmospheric nitrogen and release nutrient-rich litter, but their role as ecosystem engineers is commonly ascribed only to their diazotrophic nature.

Conclusions

We propose that the P-mobilizing capacity of Proteaceae on young soils, which contain an abundance of P, but where P is poorly available, in combination with inefficient nutrient remobilization from senescing leaves allows these species to function as ecosystem engineers. We suggest that diazotrophic species that colonize young soils with strong P-sorption potential should be considered for their positive effect on P availability, as well as their widely accepted role in nitrogen fixation. Their P-mobilizing activity possibly also enhances their nitrogen-fixing capacity. These diazotrophic species may therefore facilitate the establishment and growth of species with less-efficient P-uptake strategies on more-developed soils with low P availability through similar mechanisms. We argue that the significance of cluster roots and high carboxylate exudation in the development of young ecosystems is probably far more important than has been envisaged thus far.     

High abundance of non-mycorrhizal plant species in severely phosphorus-impoverished Brazilian <Emphasis Type="Italic">campos rupestres</Emphasis>

Background and aims

We sought to describe the species and functional composition of Brazilian campos rupestres plant communities on severely nutrient-impoverished white sands, to test hypotheses relating plant communities and physiological adaptations to infertile soils. Based on recently-published information on a south-western Australian dune chronosequence, we hypothesised that campos rupestres plant communities would similarly contain a relatively large proportion of non-mycorrhizal species, because of the phosphorus-(P) impoverished nature of the soils. We also sought to test the hypothesis that many of these non-mycorrhizal species have high leaf manganese (Mn) concentrations as a consequence of carboxylate exudation to mobilise soil P.

Methods

We conducted flora surveys and quantified mycorrhizal status and foliar Mn concentrations in field sites with strongly-weathered sandy soils. Rhizosphere carboxylates were collected from glasshouse-grown plants to assess a potential correlation of carboxylates and leaf Mn concentrations.

Results

Soils were depleted of all major plant nutrients. Non-mycorrhizal plants were abundant in most field sites (mean relative cover = 48%). Vellozia species were dominant aboveground; belowground, roots were colonised more by dark septate endophytic fungi than by mycorrhizal fungi. From the field sites, foliar Mn concentrations in non-mycorrhizal species increased with decreasing soil P concentrations, but only when soil Mn concentrations were above a minimum threshold (exchangeable [Mn] above detection limit). Across all species, however, there was no relationship of foliar Mn concentrations with soil P concentrations.

Conclusions

Our hypothesis that white-sand campos rupestres communities contain a relatively large proportion of non-mycorrhizal plants was supported. Comparison with similar ecosystems in south-western Australia suggests that plant communities on severely P-impoverished sandy soils, despite differing evolutionary histories and little overlap in plant families, follow convergent evolutionary paths towards increasing abundance of non-mycorrhizal species.

     

Mycorrhizal associations and other means of nutrition of vascular plants: understanding the global diversity of host plants by resolving conflicting information and developing reliable means of diagnosis

A comprehensive appraisal of the mycorrhizal literature provides data for 336 plant families representing 99% of flowering plants, with regard to mycorrhizas and other nutritional adaptations. In total, arbuscular (AM), orchid, ectomycorrhizas (EM) and ericoid mycorrhizas and nonmycorrhizal (NM) roots occur in 74%, 9%, 2%, 1% and 6% of Angiosperm species respectively. Many families of NM plants have alternative nutritional strategies such as parasitism, carnivory, or cluster roots. The remaining angiosperms (8%) belong to families reported to have both AM and NM species. These are designated as NM-AM families here and tend to occur in habitats considered non-conducive to mycorrhizal fungi, such as epiphytic, aquatic, extremely cold, dry, disturbed, or saline habitats. Estimated numbers of species in each category of mycorrhizas is presented with lists of NM and EM families. Evolutionary trends are also summarised by providing data on all clades and orders of flowering and non-flowering vascular plants on a global scale. A case study of Western Australian plants revealed that plants with specialised nutritional modes such as carnivory, cluster roots, or EM were much more diverse in this ancient landscape with infertile soils than elsewhere. Detailed information on the mycorrhizal diversity of plants presented here is linked to a website (mycorrhizas.info) to allow data to remain current. Over a century of research effort has resulted in data on mycorrhizal associations of >10,000 plant species that are of great value, but also somewhat of a liability due to conflicting information about some families and genera. It is likely that these conflicts result in part from misdiagnosis of mycorrhizal associations resulting from a lack of standardisation in criteria used to define them. Families that contain both NM and AM species provide a second major source of inconsistency, but even when these are excluded there is a ～10% apparent error rate in published lists of mycorrhizal plants. Arbuscules are linked to AM misdiagnosis since they are used less often than vesicles to recognise AM associations in roots and apparently occur sporadically in NM plants. Key issues with the diagnosis of mycorrhizal plants are discussed using the Cyperaceae as a case study. Detailed protocols designed to consistently distinguish AM from endophytic Glomeromycotan Fungus Colonisation (GFC) are provided. This review aims to stimulate debate and provide advice to researchers delving into root biology.     

Above- and below-ground resource acquisition strategies determine plant species responses to nitrogen enrichment

Background and AimsKnowledge of plant resource acquisition strategies is crucial for understanding the mechanisms mediating the responses of ecosystems to external nitrogen (N) input. However, few studies have considered the joint effects of above-ground (light) and below-ground (nutrient) resource acquisition strategies in regulating plant species responses to N enrichment. Here, we quantified the effects of light and non-N nutrient acquisition capacities on species relative abundance in the case of extra N input.MethodsBased on an N-manipulation experiment in a Tibetan alpine steppe, we determined the responses of species relative abundances and light and nutrient acquisition capacities to N enrichment for two species with different resource acquisition strategies (the taller Stipa purpurea, which is colonized by arbuscular mycorrhizal fungi, and the shorter Carex stenophylloides, which has cluster roots). Structural equation models were developed to explore the relative effects of light and nutrient acquisition on species relative abundance along the N addition gradient.Key ResultsWe found that the relative abundance of taller S. purpurea increased with the improved light acquisition along the N addition gradient. In contrast, the shorter C. stenophylloides, with cluster roots, excelled in acquiring phosphorus (P) so as to elevate its leaf P concentration under N enrichment by producing large amounts of carboxylate exudates that mobilized moderately labile and recalcitrant soil P forms. The increased leaf P concentration of C. stenophylloides enhanced its light use efficiency and promoted its relative abundance even in the shade of taller competitors.ConclusionsOur findings highlight that the combined effects of above-ground (light) and below-ground (nutrient) resources rather than light alone (the prevailing perspective) determine the responses of grassland community structure to N enrichment.     

Root structure and functioning for efficient acquisition of phosphorus: Matching morphological and physiological traits

BACKGROUND: Global phosphorus (P) reserves are being depleted, with half-depletion predicted to occur between 2040 and 2060. Most of the P applied in fertilizers may be sorbed by soil, and not be available for plants lacking specific adaptations. On the severely P-impoverished soils of south-western Australia and the Cape region in South Africa, non-mycorrhizal species exhibit highly effective adaptations to acquire P. A wide range of these non-mycorrhizal species, belonging to two monocotyledonous and eight dicotyledonous families, produce root clusters. Non-mycorrhizal species with root clusters appear to be particularly effective at accessing P when its availability is extremely low. SCOPE: There is a need to develop crops that are highly effective at acquiring inorganic P (Pi) from P-sorbing soils. Traits such as those found in non-mycorrhizal root-cluster-bearing species in Australia, South Africa and other P-impoverished environments are highly desirable for future crops. Root clusters combine a specialized structure with a specialized metabolism. Native species with such traits could be domesticated or crossed with existing crop species. An alternative approach would be to develop future crops with root clusters based on knowledge of the genes involved in development and functioning of root clusters. CONCLUSIONS: Root clusters offer enormous potential for future research of both a fundamental and a strategic nature. New discoveries of the development and functioning of root clusters in both monocotyledonous and dicotyledonous families are essential to produce new crops with superior P-acquisition traits.     

Characterisation of arbuscular mycorrhizal fungi colonisation in cluster roots of shape Hakea verrucosa F. Muell (Proteaceae), and its effect on growth and nutrient acquisition in ultramafic soil

Ultramafic soils at Bandalup Hill (Western Australia) are characterised by high concentrations of Ni and low levels of P. Amongst the plant species that can sustain such hostile conditions, Hakea verrucosa F. Muell from a non-mycorrhizal family (Proteaceae) would be expected to rely on cluster roots to access P. However, the acidification of ultramafic soils by cluster roots might increase the dissolution of soil Ni, and therefore its availability to plants. Symbiosis with mycorrhizal fungi, on the other hand, might help to reduce the uptake of Ni by H. verrucosa. Therefore, the aim of this study was to investigate the mycorrhizal status of H. verrucosa, and assess any contribution from mycorrhizal fungi to its growth and nutrient status. Seedlings of H. verrucosa were first grown in undisturbed ultramafic soil cores from Bandalup Hill for 8 weeks to assess the presence of mycorrhizal fungi in their roots. In a second experiment, H. verrucosa seedlings were grown in the same ultramafic soil that was either steamed or left untreated. Seedlings were inoculated with an arbuscular mycorrhizal (AM) fungal consortium from Bandalup Hill. Fungal hyphae, vesicles, as well as intracellular arbuscules and hyphal coils were observed in the cluster roots of H. verrucosa in both experiments. In the first experiment, 57% of the root length was colonized by AM fungi. Seedlings had high (between 1.4 and 1.9) shoot to root ratios and their roots had very few root hairs, despite growing in P-deficient soil. Steaming of the ultramafic soil increased the growth of seedlings and their nutrient uptake. Inoculation with AM fungi reduced the seedling growth in steamed ultramafic soil; however, it increased their shoot P and K concentration and also the shoot K content. The shoot Ni concentration of seedlings was not affected by the presence of AM fungi.     

Effects of liming and mycorrhizal colonization on soil phosphate depletion and phosphate uptake by maize (Zea mays L.) and soybean (Glycine max L.) grown in two tropical acid soils

The effects of liming and inoculation with the arbuscular mycorrhizal fungus, Glomus intraradices Schenck and Smith on the uptake of phosphate (P) by maize (Zea mays L.) and soybean (Glycine max [L.] Merr.) and on depletion of inorganic phosphate fractions in rhizosphere soil (Al-P, Fe-P, and Ca-P) were studied in flat plastic containers using two acid soils, an Oxisol and an Ultisol, from Indonesia. The bulk soil pH was adjusted in both soils to 4.7, 5.6, and 6.4 by liming with different amounts of CaCO₃.In both soils, liming increased shoot dry weight, total root length, and mycorrhizal colonization of roots in the two plant species. Mycorrhizal inoculation significantly increased root dry weight in some cases, but much more markedly increased shoot dry weight and P concentration in shoot and roots, and also the calculated P uptake per unit root length. In the rhizosphere soil of mycorrhizal and non-mycorrhizal plants, the depletion of Al-P, Fe-P, and Ca-P depended in some cases on the soil pH. At all pH levels, the extent of P depletion in the rhizosphere soil was greater in mycorrhizal than in non-mycorrhizal plants. Despite these quantitative differences in exploitation of soil P, mycorrhizal roots used the same inorganic P sources as non-mycorrhizal roots. These results do not suggest that mycorrhizal roots have specific properties for P solubilization. Rather, the efficient P uptake from soil solution by the roots determines the effectiveness of the use of the different soil P sources. The results indicate also that both liming and mycorrhizal colonization are important for enhancing P uptake and plant growth in tropical acid soils.     

Formation of cluster roots and mycorrhizal status of Comptonia peregrina and Myrica pensylvanica in Maine, USA

Extent of cluster (proteoid) root formation in the field was examined in relation to availability of P and successional status of the site in two actinorhizal, N₂-fixing shrubs. Comptonia peregrina (L.) Coult. and Myrica pensylvanica Loisel. In C. peregrina cluster roots were present at all 13 sites and comprised between 0.2 and 19% of total fine root dry weight. Cluster root formation was most extensive at recently disturbed sites and was negatively correlated with cover of associated woody species (r =−0.85), litter depth (r =−0.81) and available (extractable) soil P (r =−0.75). In M. pensylvanica cluster roots were present at all 11 sites and comprised between 6 and 20% of total fine root dry weight. On mineral soils (n = 6), extent of cluster root formation was negatively correlated (r =−0.86) with cover of associated woody species and with litter depth (r =−0.78; P = 0.07). Comptonia peregrina and M. pensylvanica lack functional mycorrhizae at these sites because only intracellular infections were found and these lacked arbuscules. None of 22 non-N₂-fixing, dominant woody species associated with C. peregrina and M. pensylvanica formed cluster roots. Cluster roots probably allow C. peregrina and M. pensylvanica to obtain sufficient P from nutrient-poor soils in the absence of functional mycorrhizae. Extensive cluster root formation on disturbed, early successional sites may aid these species in colonizing these sites because they have to form only one symbiosis (with Frankia ) and not two (with Frankia and a mycorrhizal fungus).     

常绿阔叶林外生、内生菌根树种细根化学计量学性状对N添加的响应

为揭示不同菌根类型树种细根化学计量学性状对N添加的塑性响应,在福建省建瓯市万木林自然保护区常绿阔叶林内选择外生菌根树种罗浮栲(Castanopsis faberi)和内生菌根树种木荷(Schima superba)为研究对象,采用根袋法开展N添加试验,细根在根袋内生长半年后测定化学计量学指标(C、N、P、C/N、N/P、C/P)。结果表明:根序对细根化学计量学性状有显著影响,随着根序的增加,罗浮栲与木荷细根C浓度、C/N、C/P明显增加,N浓度与P浓度明显下降。N添加对细根C、N浓度均有极显著的促进作用,但对细根P浓度影响不显著,从而导致细根C/N维持稳定,但N/P、C/P升高,细根受P限制增加。细根化学计量学性状对N添加的塑性响应在不同序级间以及在外生菌根树种罗浮栲和内生菌根树种木荷之间均无显著差异。结论表明,研究所选内生、外生菌根树种细根化学计量学性状对N添加具有基本相似的响应。     

The biology of mycorrhiza in the Ericaceae. XX. Plant and mycorrhizal necromass as nitrogenous substrates for the ericoid mycorrhizal fungus Hymenoscyphus ericae and its host

In the 'F' horizons of acid mor-humus soils of heathland ecosystems, mycorrhizal roots of the dominant ericaceous species form a large fraction of the soil biomass. Rapid turnover of these roots provides the potential for recycling of substantial amounts of nitrogen contained in their fungal and plant components. Here, we first determine the amount of N in the biomass of ericoid roots growing in heathland and show it to constitute a large proportion of total soil N. In order to assess the accessibility of this N to ericaceous plants, experiments were then conducted using aseptically produced shoot and root necromass of Vaccinium macrocarpon Ait., the roots being grown with or without mycorrhizal colonization. These materials were provided as sole nitrogenous substrates in growth experiments using the ericoid mycorrhizal fungus Hymenoscyphus ericae (Read) Korf & Kernan in pure culture and V. macrocarpon in the mycorrhizal (M) or non-mycorrhizal (NM) condition as test organisms. The experiments were designed to test the hypothesis that the N contained in these substrates can be mobilized by the mycorrhizal endophyte. The ability of the endophyte to utilize the substrates was determined by measuring fungal yields and by assessing the presence of its extra-cellular protease and chitinase enzymes. Transfer of N to the host by the endophyte was determined through measurements of plant yield and tissue N contents. H. ericae produced a significantly greater yield on shoot and mycorrhizal root necromass than on non-mycorrhizal root necromass. The extra-cellular enzymes protease and chitinase were produced by the fungus when grown on the M root necromass. The fungus also transferred N to the host plant, up to 76% of N contained in the substrate being found in M plants whereas less than 5% was present in their NM counterparts.     

大兴安岭典型森林沼泽植物叶片和细根碳氮磷化学计量特征

探究大兴安岭典型森林沼泽不同植物叶片和细根生态化学计量特征,能够为进一步认识高纬度气候敏感生态系统养分利用策略和物质循环过程提供依据。对大兴安岭地区兴安落叶松-苔草、兴安落叶松-笃斯越桔-藓类和兴安落叶松-杜香-泥炭藓3种典型森林沼泽19种优势和亚优势维管植物叶片和细根碳氮磷计量特征(C∶N∶P)进行比较,分析不同森林沼泽类型、植物生长型和菌根类型叶片和细根C∶N∶P差异,通过标准化主轴回归分析叶片与细根C∶N∶P的关系。结果表明: 叶片C∶N∶P在种间水平具有最大的变异(42.5%～84.6%),且叶片和细根种间变异大小均为N∶P>C∶N>C∶P。土壤养分和水分含量较高的兴安落叶松-苔草沼泽叶片与细根C∶N和C∶P值较低,且3种森林沼泽植物叶片和细根N∶P均小于10,受N限制。草本植物叶片C∶P和细根C∶N、C∶P显著低于木本植物。外生菌根和杜鹃花类菌根植物叶片和细根C∶N和C∶P高于丛枝菌根和无菌根植物,且杜鹃花类菌根植物叶片和细根C∶P显著高于外生菌根植物。不同森林沼泽、生长型、菌根类型植物叶片和细根C∶N和C∶P差异明显,而N∶P相对稳定。森林沼泽植物叶片与细根C∶N、C∶P和N∶P呈线性正相关,植物地上与地下部分在生态化学计量特征上存在协同。     

Cluster Roots of Leucadendron laureolum (Proteaceae) and Lupinus albus (Fabaceae) Take Up Glycine Intact: An Adaptive Strategy to Low Mineral Nitrogen in Soils?

BACKGROUND AND AIMS: South African soils are not only low in phosphorus (P) but most nitrogen (N) is in organic form, and soil amino acid concentrations can reach 2.6 g kg(-1) soil. The Proteaceae (a main component of the South African Fynbos vegetation) and some Fabaceae produce cluster roots in response to low soil phosphorus. The ability of these roots to acquire the amino acid glycine (Gly) was assessed. METHODS: Uptake of organic N as 13C-15N-Gly was determined in cluster roots and non-cluster roots of Leucadendron laureolum (Proteaceae) and Lupinus albus (Fabaceae) in hydroponic culture, taking account of respiratory loss of 13CO2. KEY RESULTS: Both plant species acquired doubly labelled (intact) Gly, and respiratory losses of 13CO2 were small. Lupin (but not leucadendron) acquired more intact Gly when cluster roots were supplied with 13C-15N-Gly than when non-cluster roots were supplied. After treatment with labelled Gly (13C : 15N ratio = 1), lupin cluster roots had a 13C : 15N ratio of about 0.85 compared with 0.59 in labelled non-cluster roots. Rates of uptake of label from Gly did not differ between cluster and non-cluster roots of either species. The ratio of C : N and 13C : 15N in the plant increased in the order: labelled roots < rest of the root < shoot in both species, owing to an increasing proportion of 13C translocation. CONCLUSIONS: Cluster roots of lupin specifically acquired more intact Gly than non-cluster roots, whereas Gly uptake by the cluster and non-cluster roots of leucadendron was comparable. The uptake capacities of cluster roots are discussed in relation to spatial and morphological characteristics in the natural environment.     

The effect of phosphorus on cluster-root formation and functioning of Embothrium coccineum (R. et J. Forst.)

Background and aims

Embothrium coccineum (R. et J. Forst.) is a Proteaceae species from the southern part of South America. South-central Chilean soils are younger and contain more phosphorus (P) than soils in Australia and South Africa, where Proteaceae are common. Phosphorus deficiency is the main factor promoting cluster-root formation in Proteaceae. It is not known, however, whether this also applies to E. coccineum, which grows on soils with higher P content.

Methods

Four-month-old seedlings were grown for 4 weeks in hydroponic cultures with 1 μM P or 50 μM P. The number of cluster roots, relative height increment, biomass distribution, cluster root/total root biomass ratio, foliar P concentration, root acid phosphatase activity and root carboxylate-exudation rates were determined.

Results

Seedlings growing at 50 μM P showed a 10?, 1.3? and 3.3-fold greater increase in relative height, total dry mass and foliar P concentration, respectively, compared with those grown at1 μM P. However, seedlings grown at 1 μM P showed a 5?, 16?, 1.7? and 1.3-fold greater number of cluster roots, cluster root/total root biomass ratio, phosphatase activity and total carboxylate exudation, respectively, as compared with those grown at 50 μM P.

Conclusions

A low P supply promotes the initiation, growth and metabolic activity of cluster roots which is in accordance with reports on Proteaceae species occurring in ancient and highly weathered soils.     

Arbuscular mycorrhizal fungi enhance P uptake and alter plant morphology in the invasive plant Microstegium vimineum

Invasive plant species can interact with native soil microbes in ways that change how they use nutrients and allocate biomass. To examine whether Microstegium vimineum form symbiotic associations with arbuscular mycorrhizal fungi (AMF) and whether AMF mediate nutrient acquisition and growth of the plant, we conducted a field survey in Raleigh, NC and Hangzhou, China and two experiments in growth chambers. This is the first report that M. vimineum is mycorrhizal, with colonization rates of 47 and 21 % in its native and invaded range, respectively. In the growth chamber, addition of an AMF inoculum mixture significantly promoted M. vimineum biomass accumulation in both field and sterilized soils, particularly after 64 days of growth. Arbuscular mycorrhizal fungi also increased plant phosphorous (P) uptake but did not consistently affect total plant nitrogen (N) acquisition, leading to decreases in plant N:P ratios. More interestingly, AMF significantly altered plant morphology, increasing the number of stolons and aerial roots per individual (59 and 723 %), aerial roots per gram aboveground biomass (374 %) and aerial roots per stolon (404 %). Our results suggest that mycorrhizal enhancement of plant growth by stimulating tillering may serve as another mechanism by which M. vimineum can quickly take over new territory. Future studies on invasive plant-microbial interactions are needed to understand the mechanisms through which microbes contribute to the competitive ability of invasive plants.     

Dinitrogen-fixing Acacia species from phosphorus-impoverished soils resorb leaf phosphorus efficiently

Nitrogen (N) and phosphorus (P) resorption from senescing leaves were studied, and the contribution of N and P cycling through litterfall to soil nutrient patchiness was investigated for four Acacia species in the Great Sandy Desert in north-western Australia. N and P concentrations of mature and recently shed leaves were analysed and compared; soils under the canopies of the shrubs and soils in gaps (open areas) between the shrubs were also analysed and compared for N and P concentrations. Mature leaf P concentrations of the plants were considerably lower than the global average values, and N : P ratios of mature leaves were high. Plants derived 0-75% of their leaf N from symbiotic N(2)-fixation. N-resorption efficiency was between 0 and 43%, and P-resorption efficiency was between 32 and 79%; all plants were more efficient at P resorption than at N resorption, and litter N : P ratios were significantly higher than mature leaf N : P ratios. Soils of the study sites were P-impoverished. Total soil N and P concentrations were higher under the canopy than in gaps, but bicarbonate-extractable P concentration was higher in gaps. Nutrient cycling through litterfall results in soil nutrient patchiness and forms 'islands of fertility' under the canopies of the shrubs.     

亚热带典型树种根毛特征及其与共生真菌的关系

根毛和共生真菌增加了吸收面积,提高了植物获取磷等土壤资源的能力。由于野外原位观测根表微观结构较为困难,吸收细根、根毛、共生真菌如何相互作用并适应土壤资源供应,缺乏相应的数据和理论。该研究以受磷限制的亚热带森林为对象,选取了21种典型树种,定量了根毛存在情况、属性变异,分析了根毛形态特征与共生真菌侵染率、吸收细根功能属性之间的关系,探讨了根表结构对低磷土壤的响应和适应格局。结果表明:1)在亚热带森林根毛不是普遍存在的, 21个树种中仅发现7个树种存有根毛, 4个为丛枝菌根(AM)树种, 3个为外生菌根(ECM)树种。其中,马尾松(Pinus massoniana)根毛出现率最高,为86%;2)菌根类型是理解根-根毛-共生真菌关系的关键,AM树种根毛密度与共生真菌侵染率正相关,但ECM树种根毛直径与共生真菌侵染率负相关; 3) AM树种根毛长度和根毛直径、ECM树种根毛出现率与土壤有效磷含量呈负相关关系。该研究揭示了不同菌根类型树种根毛-共生真菌-根属性的格局及相互作用,为精细理解养分获取策略奠定了基础。     

土壤解磷微生物促进植物磷素吸收策略研究进展

随着全球磷肥需求增加和磷矿资源储量短缺矛盾逐渐加重,土壤解磷微生物在促进磷循环方面的重要性日益突显,因此有必要对其种类和促进植物磷素吸收策略进行全面梳理总结。荟萃分析了土壤解磷微生物种类并构建了系统进化树,重点论述了土壤解磷微生物促进植物磷素吸收主要策略。土壤解磷微生物主要通过矿化和溶解作用直接活化难溶性磷,但其也能与植物根系互作间接活化磷素。其间接途径主要包括解磷微生物与根系分泌物和丛枝菌根真菌互作,它们通过碳磷交换间接活化磷素;其次包括一些解磷微生物可以通过固氮作用使植物生长受到磷的限制,从而调节植物磷酸盐转运系统间接活化土壤磷;解磷微生物也能通过分泌植物激素和生物防治剂促进植物根系生长间接促进植物磷素吸收。解磷微生物还能通过磷素固定减少磷流失,也可以通过加速自身磷素周转促进植物磷素吸收。对完善和发展解磷微生物主导的土壤磷循环和植物磷素吸收利用理论体系具有重要意义。     

Nutrient conservation strategies in native Andean‐Patagonian forests

Abstract. Nutrient conservation in vegetation affects rates of litter decomposition and soil nutrient availability. Although resorption has been traditionally considered one of the most important plant strategies to conserve nutrients in temperate forests, long leaf life‐span and low nutrient requirements have been postulated as better indicators. We aimed at identifying nutrient conservation strategies within characteristic functional groups of NW Patagonian forests on Andisols. We analysed C‐, N‐, P‐, K‐ and lignin‐concentrations in mature and senescent leaves of ten native woody species within the functional groups: broad‐leaved deciduous species, broad‐leaved evergreens and conifers. We also examined mycorrhizal associations in all species. Nutrient concentration in mature leaves and N‐ resorption were higher in broad‐leaved deciduous species than in the other two functional groups. Conifers had low mature leaf nutrient concentrations, low N‐resorption and high lignin/N ratios in senescent leaves. P‐ and K‐resorptions did not differ among functional groups. Broad‐leaved evergreens exhibited a species‐dependent response. Nitrogen in mature leaves was positively correlated with both N resorption and soil N‐fertility. Despite the high P‐retention capacity of Andisols, N appeared to be the more limiting nutrient, with most species being proficient in resorbing N but not P. The presence of endomycorrhizae in all conifers and the broad‐leaved evergreen Maytenus boaria, ectomycorrhizae in all Nothofagus species (four deciduous, one evergreen), and cluster roots in the broad‐leaved evergreen Lomatia hirsuta, would be possibly explaining why P is less limiting than N in these forests.