Mechanisms of Central Die-back of Reynoutria japonica in the Volcanic Desert on Mt. Fuji. A Stochastic Model Analysis of Rhizome Growth

Reynoutria japonica (Polygonaceae) is a pioneer clonal herbcolonising the volcanic desert on Mt. Fuji (height 3776 m),central Japan; establishment of secondary successional speciesoccurs only in the central die-back parts of the clonal standsofR. japonica. Clonal stands were excavated and the morphologyand growth pattern of the rhizomes were investigated with specialreference to the mechanisms of central die-back. Four morphologicalparameters, length of mother rhizomes, and number, branchingpositions and branching angle of daughter rhizomes on respectivemother rhizomes were measured or counted, and their roles inrhizome growth were examined employing a stochastic computersimulation model of the whole stand development. The simulationsclarified that, of these four parameters, the branching anglewas the most influential determinant of the whole pattern ofshoot distribution and that the central die-back was producedwhen the mean branching angle was 40° or smaller. Theseresults strongly infer that the onset of central die-back iscaused intrinsically byR. japonica itself by having the meanbranching angle of 40°. The adaptive significance of thegrowth pattern is discussed in relation to resource acquiringand habitat exploitation strategies of this species. Below-ground morphology; branching angle; central die-back; clonal plant; computer simulation; Japanese knotweed;Reynoutria japonica Houttuyn; rhizome growth pattern     

Successional changes in the soil microbial community along a vegetation development sequence in a subalpine volcanic desert on Mount Fuji, Japan

Aims

To study the relationship between vegetation development and changes in the soil microbial community during primary succession in a volcanic desert, we examined successional changes in microbial respiration, biomass, and community structure in a volcanic desert on Mount Fuji, Japan.

Methods

Soil samples were collected from six successional stages, including isolated island-like plant communities. We measured microbial respiration and performed phospholipid fatty acid (PLFA) analysis, denaturing gradient gel electrophoresis (DGGE) analysis, and community-level physiological profile (CLPP) analysis using Biolog microplates.

Results

Microbial biomass (total PLFA content) increased during plant succession and was positively correlated with soil properties including soil water and soil organic matter (SOM) contents. The microbial respiration rate per unit biomass decreased during succession. Nonmetric multidimensional scaling based on the PLFA, DGGE, and CLPP analyses showed a substantial shift in microbial community structure as a result of initial colonization by the pioneer herb Polygonum cuspidatum and subsequent colonization by Larix kaempferi into central areas of island-like communities. These shifts in microbial community structure probably reflect differences in SOM quality.

Conclusions

Microbial succession in the volcanic desert of Mt. Fuji was initially strongly affected by colonization of the pioneer herbaceous plant (P. cuspidatum) associated with substantial changes in the soil environment. Subsequent changes in vegetation, including the invasion of shrubs such as L. kaempferi, also affected the microbial community structure.     

Growth Dynamics of Shoot Height and Foliage Structure of a Rhizomatous Perennial Herb, Polygonum cuspidatum

The growth dynamics of shoot populations of Polygonum cuspidatumwere investigated at the Houei crater (approx. 2380 m abovesea level) on the south-eastern slope of Mount Fuji. At thisstudy site, a genetic individual of this species produces apopulation of shoots in the form of a patch occupying a certainground area. Generally, genetic individuals are located awayfrom each other and hence there is little interaction betweenindividuals. A large-sized individual occupying 31·2m² ground area with shoots, a medium-sized individual (5·6m²) and a small-sized individual (1·4 m²) were selectedfor this study. In each individual, growth was investigatedat the shoot level. The results were analysed based on the diffusionmodel. Early in the growing season in 1990, there was littledifference in LAI (leaf area index) and shoot density betweenthe individuals. Shoots of the small- and medium-sized individualsshowed size-independent height growth, whilst those of the larger-sizedindividual showed size-dependent height growth. Consequently,small-sized shoots of the small- and the medium-sized individualshad greater RGRs of shoot height growth than those of the largeindividual at the early stage. As a result, in the small- andmedium-sized individuals, cv (coefficient of variance) and skewnessof shoot height decreased with time. Increases in cv and skewnessof shoot height were found in the large-sized individual. Thesize-independent growth pattern of shoot height in the small-and medium-sized individuals during early growing stages isdifferent from the growth pattern of non-clonal plant species,in which plant height growth is positively size-dependent. Theexistence of a regulatory mechanism of shoot height growth issuggested for the small- and medium-sized individuals. The foliagestructure of the large-sized individual was different from thatof the medium- and small-sized individuals. The foliage structureof small- and medium-sized individuals was similar to the theoretical"optimal foliage structure" of plants. In clonal plant species,a genetic individual occupies a certain ground area with itsshoots. Therefore, "optimal foliage structure" per unit groundarea brings about maximization of photosynthetic rate for agenetic individual, which is consistent with the maximizationof fitness at the level of the individual plant.Copyright 1994,1999 Academic Press Clonal plant species, diffusion model, growth regulatory mechanism, genetic individual, maximization of fitness     

Genetic structure of Cenococcum geophilum populations in primary successional volcanic deserts on Mount Fuji as revealed by microsatellite markers

Polymorphic simple sequence repeat (SSR) markers were used to investigate the genetic structure in a Cenococcum geophilum population associated with Salix reinii in an early successional volcanic desert at Gotenba, on the south-eastern slope of Mount Fuji in Japan, and in three other populations associated with the same host at more developed sites on the mountain, one at Fujinomiya and two at Subashiri. The genotype richness of C. geophilum tended to be higher in more developed vegetation patches as well as in more developed sites, suggesting that genotype richness increased with advanced succession because new genotypes might have been introduced into these sites over time. High genotypic similarity was observed between the Gotenba and Fujinomiya populations but not between the Gotenba and Subashiri populations, suggesting that C. geophilum genotypes in Gotenba were introduced from the direction of Fujinomiya. Genotypes in the Gotenba population were clearly distinguishable into two groups. The absence of any intermediate genotype suggests the absence of frequent recombination in this C. geophilum population associated with early successional vegetation.     

Genetic structure and reproduction dynamics of Salix reinii during primary succession on Mount Fuji,as revealed by nuclear and chloroplast microsatellite analysis

The early stage of volcanic desert succession is underway on the southeastern slope of Mount Fuji. We used markers of nuclear microsatellites (simple sequence repeats; SSR) and chloroplast microsatellites (cpSSR) to investigate the population genetic structure and reproduction dynamics of Salix reinii, one of the dominant pioneer shrubs in this area. The number of S. reinii genets in a patch and the area of the largest genet within the patch increased with patch area, suggesting that both clonal growth and seedling recruitment are involved in the reproduction dynamics of S. reinii. Five polymorphic cpSSR markers were developed for S. reinii by sequencing the noncoding regions between universal sequences in the chloroplast genome. Nineteen different cpSSR haplotypes were identified, indicating that S. reinii pioneer genets were created by the long-distance dispersal of seeds originating from different mother genets around the study site, where all vegetation was destroyed during the last eruption. Furthermore, the clustered distributions of different haplotypes within each patch or plot suggested that newly colonized genets tended to be generated from seeds dispersed near the initially established mother genets. These results revealed that the establishment of the S. reinii population on the southeastern slope of Mount Fuji involved two sequential modes of seed dispersal: long-distance dispersal followed by short-distance dispersal.     

Pioneer dwarf willow may facilitate tree succession by providing late colonizers with compatible ectomycorrhizal fungi in a primary successional volcanic desert

To advance understanding of the contribution of ectomycorrhizal (ECM) fungi to tree successional processes, natural establishment patterns of secondary colonizing hosts and their ECM fungal communities were investigated with special reference to pioneer hosts. In the volcanic desert on Mount Fuji, Japan, vegetation is sparsely distributed, resembling islands in a sea of scoria. Of 509 vegetation islands in the research area, 161 contained Salix reinii (Salix), the first colonizing ECM host species. The spatial coincidence between secondary colonizing timber species and Salix was analysed, and ECM fungal communities were studied using molecular identification methods. I found 39 and 26 individuals of Betula ermanii and Larix kaempferi, respectively. Without exception, these individuals were all accompanied by Salix. The ECM fungal communities of these timber species showed high similarity to that of Salix and were dominated by generalists that were compatible with two or more plant families. In this desert, available ECM propagules are limited. Pioneer Salix may contribute to tree succession by providing adjacent late colonizers with compatible ECM fungal symbionts.     

Patch establishment and development of a clonal plant,Polygonum cuspidatum,on Mount Fuji

Microsatellite analysis was used to investigate the patch establishment and development of Polygonum cuspidatum Sieb. et Zucc, a clonal herbaceous plant that dominates the primary succession on the southeast slope of Mount Fuji. Genotypes of P. cuspidatum in 155 patches at the study site differed from each other. This indicates that P. cuspidatum patches are initially established by seed dispersed on the bare scoria field, and not by clonal rhizome extension. Genetic differentiation was estimated using the FST values between subpopulations at the study site. There was almost no genetic differentiation between subpopulations, indicating the presence of massive gene flow. The pollen fathers of seeds and maternal genets of current-year seedlings were inferred from the microsatellite allele composition by a simple exclusion method. The wide, random distribution of pollen fathers suggests that pollen dispersal occurs over a broad area. Maternal analysis showed a tendency for seed dispersal to be biased to the area nearby and down slope from the mother plants. Patch establishment under massive gene flow may result from such pollen and seed dispersal. To understand the process of patch development, aerial photographs taken from 1962 to 1999 were compared, and then genets in each of 36 patches were identified from the microsatellite genotypes of P. cuspidatum shoots. The comparison of aerial photographs showed that most of the patches enlarged each year and that some neighbouring patches combined during growth. Genet analysis demonstrated a high correlation between patch area and the area of the largest genet within it, and that new genets were recruited at the patch periphery. These findings indicate that both vegetative and sexual reproduction, i.e. rhizome extension and the establishment of new seedlings, contribute to the development of P. cuspidatum patches.     

Low Frankia inoculation potentials in primary successional sites at Mount St. Helens, Washington, USA

The ability of 23 year old volcanic soils from Mount St. Helens, USA, to nodulate actinorhizal Sitka alder (Alnus viridis ssp. sinuata) was studied by estimating Frankia Inoculation Unit density (FIU) in four types of primary successional communities using a host-plant bioassay. Rhizospheric Mycelial Inoculation Unit (MIU) density and the effects of inoculation with alder soil and phosphorus (P) on seedling growth were also examined. FIU and MIU were highest in alder thickets, lower in lupin patch and riparian zone sites, and lowest, with no FIUs, in bare areas. Seedling dry mass was higher in treatments inoculated with alder soil but not with P addition, while root:shoot mass ratios decreased with both inoculation and P. The extremely low densities of suitable symbionts in sparsely vegetated primary successional areas at Mount St. Helens may explain the surprisingly slow rate of spread of alder from nearby seed sources.     

Community structure of arbuscular mycorrhizal fungi in a primary successional volcanic desert on the southeast slope of Mount Fuji

Community structure of arbuscular mycorrhizal fungi (AMF), evaluated as spore samples and mycorrhizal roots of four herbaceous plant species, was investigated at different altitudes in a primary successional volcanic desert on Mount Fuji using molecular methods (fragment and sequence analysis of the large ribosomal subunit RNA gene). In total, 17 different AMF clades were identified, and most were members of the Glomaceae, Acaulosporaceae, and Gigasporaceae. The AMF community structures detected by spore sampling were inconsistent with those from plant roots. Of all AMF clades, six (35.3%) were detected only on the basis of spores, six (35.3%) only in roots, and five corresponded to both spores and roots (29.4%). Although an Acaulospora species was the most dominant among spores (67.1%), it accounted for only 6.8% in root samples. A species analysis of AMF communities at different altitudes demonstrated that AMF species diversity increased as altitude decreased and that the species enrichment at lower altitudes resulted from the addition of new species rather than species replacement. The inconsistencies in the species composition of spore communities with those in roots and the change in species diversity with altitude are discussed.     

Nitrogen-fixers <Emphasis Type="Italic">Alnus</Emphasis> and <Emphasis Type="Italic">Lupinus</Emphasis> influence soil characteristics but not colonization by later successional species in primary succession on Mount St. Helens

Changes to the primary successional environment caused by colonizing plants that present symbiotic associations with nitrogen-fixing bacteria were investigated at two areas on Mount St. Helens. One area was occupied by alder (Alnus viridis) thickets and old lupine (Lupinus lepidus) patches and the other area by young lupine patches and pumice barrens. Alder thicket soils had higher levels for a few soil nutrients and had greater cover by other pioneer species as compared to old lupine patches. Many soil nutrients, including nitrogen and soil organic matter, were below detection limits in old lupine patches but not in alder thicket soils. Young lupine patch soils were generally not different from barren site soils but had greater cover by other pioneer species. Below detection nitrogen and soil organic matter levels also occurred in many barren soil samples but not in young lupine patch soils. Barren soils were moister than were the other sites. The apparent increase in soil fertility has not led to invasion by later successional species, perhaps due to dry conditions or to other inhibitory factors. Seedbanks, composed of early successional species, appear to be developing in these areas.     

Nonlinear processes in seagrass colonisation explained by simple clonal growth rules

The development of single clones of the seagrass Cymodocea nodosa was analysed using a growth model based on the formation of structures limited by diffusive aggregation. The model implemented the measured clonal growth rules (i.e. rhizome elongation and branching rates, branching angle, and spacer length between consecutive shoots) and shoot mortality rate for C. nodosa at Alfacs Bay (Spain). The simulated patches increased their size nonlinearly with time displaying two different domains of growth. Young patches showed a rapid increase with time of the length of rhizome network and the number of living shoots, which depended on rhizome branching rate, and increased the radial patch size (R_g) algebraically with the number of living shoots as R_g ∝ N_s^1/Df, being D_f the fractal dimension of the patch structure. Patches older than 4 years increased the production of rhizome network and the number of living shoots much more slowly, while their radial patch size behaved as R_g ∝ N_s^0.5 resulting from an internal patch compactation. Moreover, the linear growth rate of the simulated patches changed up to 30 fold during patch development, increasing with increasing patch size until patches reached an intermediate size. The modelled patch development was found to closely reproduce the observed patch structure for the species at the Alfacs Bay (Spain). Hence, the growth of C. nodosa patches initially proceeds with a growth mode controlled by the branching pattern (branching frequency and angle) of the species, producing sparse and elongated patches. Once patches exceed 4–5 years of age and contained >500 shoots, becoming dense and circular, they shifts to a growth model typical of compact structures. These results explain previously unaccounted evidence of the emergence of nonlinear patch growth from simple clonal growth rules, and highlight the importance of branching frequency and angles as critical determinants of the space occupation rate of seagrasses and probably other clonal plants.     

Nitrogen translocation via rhizome systems in monoclonal stands ofReynoutria japonica in an oligotrophic desert on Mt Fuji: Field experiments

Mechanisms which enableReynoutria japonica, a dominant pioneer herb, to be successful in maintaining large stands in an oligotrophic volcanic desert on Mt Fuji were investigated with special reference to its nitrogen acquisition.Reynoutria japonica forms circular stands, each of which comprises only one genet. As a stand develops outwards, the number of aerial shoots per unit area decreases in the center. Shoots grow vigorously in the peripheral area where the available nitrogen from soil and precipitation (about 2.4 g m⁻² year⁻²) was much less than total nitrogen in the shoots (6.1–9.1 g m⁻²). Leaf nitrogen content per unit mass was also greater in the leaves of the peripheral shoots. When rhizomes extending radially from the center to the periphery were severed, the dry mass of shoots in the periphery diminished by 75% on a ground area basis. In the periphery, leaf nitrogen content also reduced significantly and no flowers were produced. When fertilizer was applied to the peripheral shoots with severed rhizomes, neither growth, survival nor flower production of the shoots was significantly smaller than the control levels. In these shoots, it is also found that the nitrogen content in the youngest leaves decreased for about 1 month and then increased to above that in the control leaves. These results suggest that (i) nitrogen accumulated in the central part is translocated to peripheral shoots via rhizomes, and that long-distance translocation enables the stands to develop outwards, and (ii) aerial shoots in the periphery utilize the nitrogen translocated by rhizomes in the beginning of the growth season, whereas once the shoots have established, they begin to take up nitrogen with their own roots. Since the peripheral shoots are in sunnier environments than the shoots inside the stand, the acropetal nitrogen translocation via rhizomes will raise the production efficiency of a whole stand.     

Development of microsatellite markers in polyploid Salix reinii

Salix reinii Franch. et Savat., a crawl shrub, is one of the most dominant pioneer trees on the southeastern slope of Mt. Fuji, on which an early stage of the volcanic desert succession has been underway since the last eruption in 1707. To investigate the population genetic structure within and among patches of S. reinii, seven polymorphic microsatellite markers were developed by an enrichment procedure. The multiple banding patterns amplified by developed microsatellite markers showed the polyploidy in S. reinii growing on the southeastern slope of Mt. Fuji.     

Allometric Root/Shoot Relationships and Predicted Water Uptake for Desert Succulents

Root morphology, shoot morphology, and water uptake for Agavedeserti and Ferocactus acanthodes of various sizes were studiedusing allometric relationships (y = ax^b) and a previously developedwater uptake model. Shoot surface area increased with shootvolume with an exponent b of 0.75 for both species. Root lengthand the ground area explored by the roots increased with shootsurface area with b's of 0.72 for A. deserti and 0.92 for F.acanthodes. Various sized individuals had about the same ratioof root length to explored ground area, with higher values occurringfor A. deserti. Predicted water uptake averaged over the exploredground area was approximately constant over a 10⁴-fold rangein shoot surface area, suggesting that shoot size confers nointraspecific competitive advantage for water uptake. For theroot lengths per explored ground area observed in the field,water uptake was predicted to be 85 per cent of maximal; wateruptake could be increased by the production of more rain roots.When differences in shoot volume were accounted for by allometry,small plants had relatively less shoot surface area and relativelymore root length per shoot volume than did large plants, whichmay be important for the water relations of seedling establishment. Agave deserti, Ferocactus acanthodes, allometry, desert succulents, root distribution, root length, seedling growth, seedling establishment, shoot surface area, shoot volume, water uptake     

The influence of patch demographics on metapopulations, with particular reference to successional landscapes

The classical view of metapopulations relates the regional abundance of a species to the balance between the extinction and colonization dynamics of identical local populations. Species in successional landscapes may represent the most appropriate examples of classical metapopulations. However, Levins‐type metapopulation models do not explicitly separate population loss due to successional habitat change from other causes of extinction. A further complication is that the chance of population loss due to successional habitat change may be related to the age of a patch. I developed simple patch occupancy models to include succession and included consideration of patch age structure to address two related questions: what are the implications of changes in patch demographic rates and when is a move to a structured patch occupancy model justified? Age‐related variation in patch demography could increase or decrease the equilibrium fraction of the available habitat occupied by a species when compared to the predictions of an unstructured model. Metapopulation persistence was enhanced when the age class of patches with the highest species occupancy suffered relatively low losses to habitat succession. Conversely, when the age class of patches with the highest species occupancy also had relatively high successional loss rates, extinction thresholds were higher that would be predicted by a simple unstructured model. Hence age‐related variation in patch successional rate introduces biases into the predictions of simple unstructured models. Such biases can be detected from field surveys of the fraction of occupied and unoccupied patches in each age class. Where a bias is demonstrated, unstructured models will not be adequate for making predictions about the effects of changing parameters on metapopulation size. Thinking in successional terms emphasizes how landscapes might be managed to enhance or reduce the patch occupancy by any particular metapopulation     

准噶尔荒漠早春短命植物的光合特性及生物量分配特点

准噶尔荒漠分布的早春短命植物不仅具有十分独特的生物学特点,而且在荒漠植物群落演替、物种多样性维持及土壤改良与防治水土流失等方面具有重要的生态学价值。该文运用Li-6400开放式气体交换光合作用测定系统,对分布于准噶尔荒漠的16种早春短命植物生长盛期的净光合速率(P_n)、蒸腾速率(T_r)、水分利用效率(WUE)等特征进行了测定,并对其中7种植物与生长相关的生物量分配特征进行了分析。结果表明:1)16种植物的最大P_n、 最大T_r及WUE分别为8.07~35.96 μmol CO₂·m^-2·s^-1、3.16~29.64 mmol H₂O·m^-2·s^-1、0.54~4.26 μmol CO₂·mmol^-1H₂O;种间最大P_n与最大气孔导度(Stomatal conductance, G_s)之间存在正相关关系,其相关系数为0.77(p<0.05),线性回归斜率为26.36 μmol·mmol^-1;从光合速率对胞间CO₂浓度及光量子通量密度的响应曲线来看,这类植物的表观CO₂补偿点均在4~5 Pa之间(28~30 ℃),表观羧化效率为0.64~1.86 μmol CO₂·m^-2·s^-1·Pa^-1,表观量子效率为0.05~0.06。2)从生物量分配来看,所测植物的个体生物量为0.05~0.39 g;单株总叶面积为 3.24~51.40 cm²;单位叶面积干重为0.40~0.77 g·m^-2,根在总生物量中所占比例为5.72%~19.43%,单株叶面积比在2.92~9.00 m²·kg^-1之间。种间根所占生物量的比与对应的WUE之间的比较分析结果表明,二者之间存在显著的正相关关系,其相关系数r为0.93(p<0.01)。这些结果表明,所观测的早春短命植物具有典型的C₃植物特征,相比其它类型的荒漠植物具有较高的单位叶面积P_n、高T_r及低WUE,并且在生长发育过程中表现出很低的根/地上生物量比、较高的叶面积比和单位叶面积干重,说明它们具有相对高的生长速率,这与其生长发育节律相一致,反映了它们与准噶尔荒漠环境相适应的特点。     

Plant species associations in alpine-subnival vegetation patches in the Central Caucasus

Abstract. Patches of herbaceous vegetation found at the boundary of the alpine and subnival belts in the Central Caucasus (3000 m) were analysed for species composition, interspecific associations, as well as for relations between age, size and diversity of the patches. 34 plant species were recorded. Positive and negative associations were found among the most frequent species. A model is presented that describes the cumulation of species in patches during succession. In the final and stable successional stage, the maximum number of species within a patch was nine. Species composition varies in patches of different age, typical alpine species being more frequent in older ones. The results are interpreted as evidence for niche differentiation and selection of species composition through the 'sieve’ of interspecific relations, taking place during succession in the studied patches.     

松嫩草原水淹恢复演替过程中羊草无性系种群构件的物质生产与贮藏

在松嫩平原羊草草甸,对水淹恢复演替过程中半径为0.5、1、3、和10 m的羊草无性系种群斑块分蘖株的物质生产和根茎的物质贮藏进行了研究.结果表明,0.5、1、3、 m斑块各圈和10 m中间及边缘根茎的物质贮藏以1龄级、不同斑块各圈分蘖株的物质生产以1、2龄级对种群的贡献最大,其物质贮藏和生产所占比例均逐圈增加.整体上,年轻龄级分蘖株和根茎具有旺盛的物质生产力和贮藏力.无性系斑块由中心至外围的物质生产和贮藏规律与斑块扩展趋势具有一致性.0.5～10 m斑块分蘖株物质生产力和根茎贮藏力的变异以最外圈最大.在水淹羊草草甸的恢复过程中,不同大小羊草无性系斑块种群构件的物质生产和贮藏策略为种群不断扩展和群落进展演替奠定了物质基础.     

Growth Consequences of Soil Nutrient Heterogeneity for two Old-field Herbs,Ambrosia artemisiifoliaandPhytolacca americana, Grown Individually and in Combination

Plant species can respond to small scale soil nutrient heterogeneityby proliferating roots or increasing nutrient uptake kineticsin nutrient-rich patches. Because root response to heterogeneitydiffers among species, it has been suggested that the distributionof soil resources could influence the outcome of interspecificcompetition. However, studies testing how plants respond toheterogeneity in the presence of neighbours are lacking. Inthis study, individuals of two species,Phytolacca americanaL.andAmbrosia artemisiifoliaL. were grown individually and incombination in soils with either a homogeneous or heterogeneousnutrient distribution. Above-ground biomass of individuallygrown plants of both species was greater when fertilizer waslocated in a single patch than when the same amount of fertilizerwas distributed evenly throughout the soil. Additionally, bothspecies proliferated roots in high-nutrient patches.A. artemisiifoliaexhibitedlarger root:shoot ratios, increased nitrogen depletion fromnutrient patches, and a higher growth rate thanP. americana,suggestingA. artemisiifoliais better suited to find and rapidlyexploit nutrient patches. In contrast to individually grownplants, soil nutrient distribution had no effect on final above-groundplant biomass for either species when grown with neighbours,even though roots were still concentrated in high nutrient patches.This study demonstrates that increased growth of isolated plantsas a consequence of localized soil nutrients is not necessarilyan indication that heterogeneity will affect interspecific encounters.In fact, despite a significant below-ground response, soil nutrientheterogeneity was inconsequential to above-ground performancewhen plants were grown with neighbours.Copyright 1999 Annalsof Botany Company Phytolacca americana, pokeweed,Ambrosia artemisiifolia, ragweed, nutrient heterogeneity, root proliferation, plasticity, foraging, nutrient patches.     

Germination and infectivity of ectomycorrhizal fungal spores in relation to their ecological traits during primary succession

The spores of ectomycorrhizal fungi (EMF) play critical roles in the population and community development of EMF. Here, the germination and infectivity of EMF spores are examined with reference to the ecological traits of the EMF species. Spores were collected from 12 EMF species, whose successional patterns have been studied in the volcanic desert on Mount Fuji, Japan. Spore germination experiments were conducted with host plants (Salix reinii), with nonhost plants (Polygonum cuspidatum), and without plants. The mycorrhizal formation ability of spores was also examined in seven EMF using spore inoculation experiments. To determine the effects of the spore preservation period, both experiments were repeated up to 1 yr after spore collection. Spore germination was very low in the absence of host plants. In the presence of hosts, even 30 d after spore collection, spore germination was significantly enhanced in all pioneer EMF (c. 20%) but less so in late-stage EMF (< 5%), except in Hebeloma species. Mycorrhizal formation from spores was also greater in pioneer EMF but was significantly reduced by 1 yr of spore preservation. High spore germination and infectivity of pioneer EMF should enable these species to colonize disturbed and isolated areas in accordance with their ecological traits.