Phosphorus and intraspecific density alter plant responses to arbuscular mycorrhizas

We investigated how phosphorus availability, intraspecific density, and their interaction affect plant responses to arbuscular mycorrhizas. Four facultatively mycotrophic species: chile, cilantro, tomato, and corn were examined separately in pot experiments that employed a tropical phosphorus-immobilizing soil. Each experiment comprised nine soluble phosphorus additions, two levels of intraspecific plant density, and inoculation with arbuscular mycorrhizal fungi or not. High phosphorus signi- ficantly diminished mycorrhizal colonization of corn, cilantro, and tomato, but not chile, which was highly variably colonized. Corn roots were colonized by other root-inhabiting fungi, and mycorrhizas significantly reduced colonization by these potential root parasites. High phosphorus significantly increased relative growth rates (RGR) of all species, and high density significantly reduced RGR of cilantro, tomato, and corn. Chile showed little growth at any but the highest phosphorus additions, and consequently had no RGR response to density or mycorrhizas. Mycorrhiza inoculation caused transient depression of corn growth during the first month, but mycorrhizas increased corn RGR during the second month of growth. Both RGR and dry weights at harvest, cilantro, tomato, and corn benefited from mycorrhizas at low phosphorus availability, but this benefit diminished or changed to disadvantage as phosphorus availability increased. At low phosphorus availability, high density increased the dry weight of mycorrhizal cilantro and thereby amplified the benefit of mycorrhizas. At high phosphorus availability, increased density diminished the effects of mycorrhizas on dry weight, reducing mycorrhiza benefit to tomato and chile and reducing mycorrhiza detriment to cilantro. This study demonstrates that for three of the four plant species examined, phosphorus availability, intraspecific density, and their interaction significantly modify plant responses to arbuscular mycorrhizas.     

The screening and selection of inoculant arbuscular-mycorrhizal and ectomycorrhizal fungi

The initiation of a programme of screening and selection of arbuscular-mycorrhizal fungi (AM fungi) and ectomycorrhizal fungi (ECM fungi) for use as inoculant fungi in agriculture, horticulture of forestry will depend on whether inoculation is more appropriate as a management option than manipulation of the indigenous mycorrhizal populations. The greatest immediate potential for the successful use of mycorrhizal inoculants will be in soils and growth media where phosphorus (P) limits plant growth and where the indigenous mycorrhizal fungi are either ineffective at increasing P uptake by the plant or their numbers have been drastically reduced by human influence or natural disturbance. In recent investigations, however, additional benefits to the plant following colonization of roots by effective AM or ECM fungi have been demonstrated. These additional benefits of an effective mycorrhizal association have necessitated a re-evaluation of the optimum screening procedures for isolates of AM and ECM fungi. Both current methodologies and suggestions for alternative approaches to the screening of AM and ECM fungi are discussed in this paper. The problems inherent in choosing suitable experimental conditions to compare different isolates at the screening stage are also addressed. The review also stresses the importance of continued monitoring of introduced mycorrhizal fungi to learn more about their longer-term ecological role, particularly within reforestation or revegetation studies.     

Effects of mycorrhizal symbiosis on tallgrass prairie plant–herbivore interactions

Complex relationships occur among plants, mycorrhizal fungi, and herbivores. By altering plant nutrient status, mycorrhizas may alter herbivory or plant tolerance to herbivory via compensatory regrowth. We examined these interactions by assessing grasshopper preference and plant growth and fungal colonization responses to herbivory under mycorrhizal and non‐mycorrhizal conditions within tallgrass prairie microcosms. Mycorrhizal symbiosis increased plant regrowth following defoliation, and some strongly mycotrophic plant species showed overcompensation in response to herbivory when they were mycorrhizal. Although grasshoppers spent more time on mycorrhizal plants, herbivory intensity did not differ between mycorrhizal and non‐mycorrhizal plants. Aboveground herbivory by grasshoppers significantly increased mycorrhizal fungal colonization of plant roots. Thus mycorrhizas may greatly benefit plants subjected to herbivory by stimulating compensatory growth, and herbivores, in turn, may increase the development of the symbiosis. Our results also indicate strong interspecific differences among tallgrass prairie plant species in their responses to the interaction of aboveground herbivores and mycorrhizal symbionts.     

Plant responsiveness to mycorrhizas differs from dependence upon mycorrhizas

Soil phosphorus response curves of plants with and without mycorrhizas reflect two different, but complementary, phenomena. The first, plant responsiveness to mycorrhizas, is represented by the difference in growth between plants with and without mycorrhizas at any designated level of phosphorus availability. This is also a measure of mycorrhizal fungus effectiveness. The second, the lowest level of phosphorus availability at which plants can grow without mycorrhizas, is here termed dependence upon mycorrhizas. The latter definition differs from conventional usage which fails to distinguish dependence from responsiveness. Sigmoid curves generated by the three-parameter, logistic equation generally can model the responses of plants to mycorrhizas and phosphorus addition and can be used to assess responsiveness, effectiveness, and dependence. Such curves reveal that plant responsiveness or fungus effectiveness determined at a single level of phosphorus availability may be misleading when used to compare different host species' intrinsic capacities to respond to different mycorrhizal fungus species. Instead, the same relative position should be evaluated among phosphorus response curves for different species combinations. Dependence of a plant species known to benefit from mycorrhizas can be assessed with reference to only the phosphorus response curve of plants without mycorrhizas. Dependence is a constitutive property of plant species that can be used to classify them as facultatively or obligately mycotrophic. Dependence is a plant attribute upon which natural selection can act, but responsiveness and effectiveness cannot be selected directly because they are emergent properties of the interaction between plant and fungus species.     

Mycorrhizas influence functional traits of two tallgrass prairie species

Over the past decade, functional traits that influence plant performance and thus, population, community, and ecosystem biology have garnered increasing attention. Generally lacking, however, has been consideration of how ubiquitous arbuscular mycorrhizas influence plant allometric and stoichiometric functional traits. We assessed how plant dependence on and responsiveness to mycorrhizas influence plant functional traits of a warm‐season, C4 grass, Andropogon gerardii Vitman, and the contrasting, cool‐season, C3 grass, Elymus canadensis L. We grew both host species with and without inoculation with mycorrhizal fungi, across a broad gradient of soil phosphorus availabilities. Both host species were facultatively mycotrophic, able to grow without mycorrhizas at high soil phosphorus availability. A. gerardii was most dependent upon mycorrhizas and E. canadensis was weakly dependent, but highly responsive to mycorrhizas. The high dependence of A. gerardii on mycorrhizas resulted in higher tissue P and N concentrations of inoculated than noninoculated plants. When not inoculated, E. canadensis was able to take up both P and N in similar amounts to inoculated plants because of its weak dependence on mycorrhizas for nutrient uptake and its pronounced ability to change root‐to‐shoot ratios. Unlike other highly dependent species, A. gerardii had a high root‐to‐shoot ratio and was able to suppress colonization by mycorrhizal fungi at high soil fertilities. E. canadensis, however, was unable to suppress colonization and had a lower root‐to shoot ratio than A. gerardii. The mycorrhiza‐related functional traits of both host species likely influence their performance in nature: both species attained the maximum responsiveness from mycorrhizas at soil phosphorus availabilities similar to those of tallgrass prairies. Dependence upon mycorrhizas affects performance in the absence of mycorrhizas. Responsiveness to mycorrhizal fungi is also a function of the environment and can be influenced by both mycorrhizal fungus species and soil fertility.     

Transport processes at the plant-fungus interface in mycorrhizal associations: physiological studies

The roots of most plants form symbiotic associations with mycorrhizal fungi. The net flux of nutrients, particularly phosphorus (P), from the soil into the plant is greater in mycorrhizal than in comparable non-mycorrhizal plants. However despite the widespread occurrence of mycorrhizal associations the processes controlling the transfer of solutes between the symbionts are poorly understood. To understand the mechanisms regulating the transfer of solutes information about conditions at the interface between plant and fungus is needed.Measurements of apoplastic and intracellular electrical potential difference in leek roots colonised by mycorrhizal fungi and estimates of cytosolic pH in fungal hyphae are presented. These and the implications for plant/fungal mineral nutrition in vesicular-arbuscular mycorrhizas are discussed.     

Mycorrhizas and soil structure

In addition to their well-recognized roles in plant nutrition and communities, mycorrhizas can influence the key ecosystem process of soil aggregation. Here we review the contribution of mycorrhizas, mostly focused on arbuscular mycorrhizal fungi (AMF), to soil structure at various hierarchical levels: plant community; individual root; and the soil mycelium. There are a suite of mechanisms by which mycorrhizal fungi can influence soil aggregation at each of these various scales. By extension of these mechanisms to the question of fungal diversity, it is recognized that different species or communities of fungi can promote soil aggregation to different degrees. We argue that soil aggregation should be included in a more complete 'multifunctional' perspective of mycorrhizal ecology, and that in-depth understanding of mycorrhizas/soil process relationships will require analyses emphasizing feedbacks between soil structure and mycorrhizas, rather than a uni-directional approach simply addressing mycorrhizal effects on soils. We finish the discussion by highlighting new tools, developments and foci that will probably be crucial in further understanding mycorrhizal contributions to soil structure.     

亚热带草地中植物优势种与从属种对AM真菌的差异性生长反应

AM真菌能够影响植物生态系统的群落结构.以亚热带草地生态系统为研究对象,调查了两块草地中优势种和从属种的菌根,并在盆栽试验中比较了优势种和从属种对AM真菌的土著菌种和外源菌种Glomus mosseae的生长反应、养分吸收.结果表明,两块草地各自的优势种藿香蓟和两耳草对土著菌种的菌根依赖性分别是41.5%和77.4%,远远高于从属种莎草和毛蓼（16.0%和7.9%）;但是它们对Glomus mosseae的菌根依赖性有所变化,分别是79.6%、44.2%、74.1%和24.9%.这表明,土著菌种是优势种和从属种的形成机制之一,而外源菌种可能改变基于土著菌种而形成的植物群落结构.植株磷营养的分析结果表明,AM真菌对优势种和从属种生长的促进与对磷吸收的促进高度相关,表明AM真菌促进养分吸收是其影响植物群落结构的机制之一.     

Molecular trait indicators: moving beyond phylogeny in arbuscular mycorrhizal ecology

Arbuscular mycorrhizal (AM) fungi form symbiotic associations with the roots of most plants, thereby mediating nutrient and carbon fluxes, plant performance, and ecosystem dynamics. Although considerable effort has been expended to understand the keystone ecological position of AM symbioses, most studies have been limited in scope to recording organism occurrences and identities, as determined from morphological characters and (mainly) ribosomal sequence markers. In order to overcome these restrictions and circumvent the shortcomings of culture- and phylogeny-based approaches, we propose a shift toward plant and fungal protein-encoding genes as more immediate indicators of mycorrhizal contributions to ecological processes. A number of candidate target genes, involved in the uptake of phosphorus and nitrogen, carbon cycling, and overall metabolic activity, are proposed. We discuss the advantages and disadvantages of future protein-encoding gene marker and current (phylo-) taxonomic approaches for studying the impact of AM fungi on plant growth and ecosystem functioning. Approaches based on protein-encoding genes are expected to open opportunities to advance the mechanistic understanding of ecological roles of mycorrhizas in natural and managed ecosystems.     

Micronutrient transport in mycorrhizal symbiosis; zinc steals the show

Mycorrhizas are mutually beneficial associations between soil-borne fungi and plant roots. Mycorrhizal fungi provide their host plant with essential nutrients in exchange for sugars and/or lipids. Traditionally, transport and translocation of macronutrients, including nitrogen and phosphorus, throughout the fungal mycelium and towards the host plant are well studied. However, the regulation of nutrient exchange and their contribution in the morphogenesis and development of mycorrhizas remains unclear. In this Opinion, we argue that adding micronutrients in the current models of symbiotic transport is essential to fully understand the establishment, maintenance, and functioning of mycorrhizal associations. Homeostatic mechanisms at the cellular level and the first transport proteins involved have been recently documented for zinc (Zn) in arbuscular mycorrhizal, ectomycorrhizal, and ericoid mycorrhizal fungi. Mycorrhizal plants benefit from an improved Zn status in control conditions and are better protected when environmental Zn availability fluctuates. These recent progresses are paving the way for a better understanding of micronutrient allocation in mycorrhizas. Revising our vision on the role of micronutrients, particularly of Zn, in these interactions will allow a better use of mycorrhizal fungi in sustainable agriculture and forestry, and will increase management practices in waste land, as well as in agricultural and natural ecosystems.     

Subcellular Nutrient Element Localization and Enrichment in Ecto- and Arbuscular Mycorrhizas of Field-Grown Beech and Ash Trees Indicate Functional Differences

Mycorrhizas are the chief organ for plant mineral nutrient acquisition. In temperate, mixed forests, ash roots (Fraxinus excelsior) are colonized by arbuscular mycorrhizal fungi (AM) and beech roots (Fagus sylvatica) by ectomycorrhizal fungi (EcM). Knowledge on the functions of different mycorrhizal species that coexist in the same environment is scarce. The concentrations of nutrient elements in plant and fungal cells can inform on nutrient accessibility and interspecific differences of mycorrhizal life forms. Here, we hypothesized that mycorrhizal fungal species exhibit interspecific differences in mineral nutrient concentrations and that the differences correlate with the mineral nutrient concentrations of their associated root cells. Abundant mycorrhizal fungal species of mature beech and ash trees in a long-term undisturbed forest ecosystem were the EcM Lactarius subdulcis, Clavulina cristata and Cenococcum geophilum and the AM Glomus sp. Mineral nutrient subcellular localization and quantities of the mycorrhizas were analysed after non-aqueous sample preparation by electron dispersive X-ray transmission electron microscopy. Cenococcum geophilum contained the highest sulphur, Clavulina cristata the highest calcium levels, and Glomus, in which cations and P were generally high, exhibited the highest potassium levels. Lactarius subdulcis-associated root cells contained the highest phosphorus levels. The root cell concentrations of K, Mg and P were unrelated to those of the associated fungal structures, whereas S and Ca showed significant correlations between fungal and plant concentrations of those elements. Our results support profound interspecific differences for mineral nutrient acquisition among mycorrhizas formed by different fungal taxa. The lack of correlation between some plant and fungal nutrient element concentrations may reflect different retention of mineral nutrients in the fungal part of the symbiosis. High mineral concentrations, especially of potassium, in Glomus sp. suggest that the well-known influence of tree species on chemical soil properties may be related to their mycorrhizal associates.     

Further advances in orchid mycorrhizal research

Orchid mycorrhizas are mutualistic interactions between fungi and members of the Orchidaceae, the world’s largest plant family. The majority of the world’s orchids are photosynthetic, a small number of species are myco-heterotrophic throughout their lifetime, and recent research indicates a third mode (mixotrophy) whereby green orchids supplement their photosynthetically fixed carbon with carbon derived from their mycorrhizal fungus. Molecular identification studies of orchid-associated fungi indicate a wide range of fungi might be orchid mycobionts, show common fungal taxa across the globe and support the view that some orchids have specific fungal interactions. Confirmation of mycorrhizal status requires isolation of the fungi and restoration of functional mycorrhizas. New methods may now be used to store orchid-associated fungi and store and germinate seed, leading to more efficient culture of orchid species. However, many orchid mycorrhizas must be synthesised before conservation of these associations can be attempted in the field. Further gene expression studies of orchid mycorrhizas are needed to better understand the establishment and maintenance of the interaction. These data will add to efforts to conserve this diverse and valuable association.     

基于SCI文献分析我国菌根学研究现状和发展方向

基于美国科学情报研究所(ISI)科学引文索引(Science Citation Index, SCI)数据库, 对1989~2007年期间我国发表的菌根研究论文进行检索, 并采用文献计量方法对所获资料进行统计和趋势分析。结果表明, 1989~2007年间, 我国菌根学研究呈现不断上升趋势, 尤其是2000年之后, 增长速度明显加快。但只有5.22%的论文发表于影响因子大于5的刊物, 在研究深度上仍需进一步加强。半数以上的研究论文与丛枝菌根研究有关, 研究重点主要集中在菌根对植物的生理效应, 菌根与植物抗性     

Patterns of plant naturalization show that facultative mycorrhizal plants are more likely to succeed outside their native Eurasian ranges

The naturalization of an introduced species is a key stage during the invasion process. Therefore, identifying the traits that favor the naturalization of non-native species can help understand why some species are more successful when introduced to new regions. The ability and the requirement of a plant species to form a mutualism with mycorrhizal fungi, together with the types of associations formed may play a central role in the naturalization success of different plant species. To test the relationship between plant naturalization success and their mycorrhizal associations we analysed a database composed of mycorrhizal status and type for 1981 species, covering 155 families and 822 genera of plants from Europe and Asia, and matched it with the most comprehensive database of naturalized alien species across the world (GloNAF). In mainland regions, we found that the number of naturalized regions was highest for facultative mycorrhizal, followed by obligate mycorrhizal and lowest for non-mycorrhizal plants, suggesting that the ability of forming mycorrhizas is an advantage for introduced plants. We considered the following mycorrhizal types: arbuscular, ectomycorrhizal, ericoid and orchid mycorrhizal plants. Further, dual mycorrhizal species were those that included observations of arbuscular mycorrhizas as well as observations of ectomycorrhizas. Naturalization success (based on the number of naturalized regions) was highest for arbuscular mycorrhizal and dual mycorrhizal plants, which may be related to the low host specificity of arbuscular mycorrhizal fungi and the consequent high availability of arbuscular mycorrhizal fungal partners. However, these patterns of naturalization success were erased in islands, suggesting that the ability to form mycorrhizas may not be an advantage for establishing self-sustaining populations in isolated regions. Taken together our results show that mycorrhizal status and type play a central role in the naturalization process of introduced plants in many regions, but that their effect is modulated by other factors.     

Mycorrhizal fungal growth responds to soil characteristics,but not host plant identity,during a primary lacustrine dune succession

Soil factors and host plant identity can both affect the growth and functioning of mycorrhizal fungi. Both components change during primary succession, but it is unknown if their relative importance to mycorrhizas also changes. This research tested how soil type and host plant differences among primary successional stages determine the growth and plant effects of arbuscular mycorrhizal (AM) fungal communities. Mycorrhizal fungal community, plant identity, and soil conditions were manipulated among three stages of a lacustrine sand dune successional series in a fully factorial greenhouse experiment. Late succession AM fungi produced more arbuscules and soil hyphae when grown in late succession soils, although the community was from the same narrow phylogenetic group as those in intermediate succession. AM fungal growth did not differ between host species, and plant growth was similarly unaffected by different AM fungal communities. These results indicate that though ecological filtering and/or adaptation of AM fungi occurs during this primary dune succession, it more strongly reflects matching between fungi and soils, rather than interactions between fungi and host plants. Thus, AM fungal performance during this succession may not depend directly on the sequence of plant community succession.     

农业生态系统中的AM真菌多样性

农业生态系统复杂庞大,是由如麦田生态系统、水稻田生态系统、果园生态系统、草地生态系统、保护地生态系统等组成的一个复合生态系统。重点介绍农业生态系统中丛枝菌根(AM)和AM真菌多样性,探讨农业生态系统中调控AM真菌多样性的途径以及今后研究的动向。     

Plant–fungus competition for nitrogen erases mycorrhizal growth benefits of Andropogon gerardii under limited nitrogen supply

Considered to play an important role in plant mineral nutrition, arbuscular mycorrhizal (AM) symbiosis is a common relationship between the roots of a great majority of plant species and glomeromycotan fungi. Its effects on the plant host are highly context dependent, with the greatest benefits often observed in phosphorus (P)‐limited environments. Mycorrhizal contribution to plant nitrogen (N) nutrition is probably less important under most conditions. Moreover, inasmuch as both plant and fungi require substantial quantities of N for their growth, competition for N could potentially reduce net mycorrhizal benefits to the plant under conditions of limited N supply. Further compounded by increased belowground carbon (C) drain, the mycorrhizal costs could outweigh the benefits under severe N limitation. Using a field AM fungal community or a laboratory culture of Rhizophagus irregularis as mycorrhizal inoculants, we tested the contribution of mycorrhizal symbiosis to the growth, C allocation, and mineral nutrition of Andropogon gerardii growing in a nutrient‐poor substrate under variable N and P supplies. The plants unambiguously competed with the fungi for N when its supply was low, resulting in no or negative mycorrhizal growth and N‐uptake responses under such conditions. The field AM fungal communities manifested their potential to improve plant P nutrition only upon N fertilization, whereas the R. irregularis slightly yet significantly increased P uptake of its plant host (but not the host's growth) even without N supply. Coincident with increasing levels of root colonization by the AM fungal structures, both inoculants invariably increased nutritional and growth benefits to the host with increasing N supply. This, in turn, resulted in relieving plant P deficiency, which was persistent in non‐mycorrhizal plants across the entire range of nutrient supplies.     

Occurrence of arbuscular mycorrhizal fungi in a phosphorus-poor wetland and mycorrhizal response to phosphorus fertilization

The presence of arbuscular mycorrhizas in fens has received little attention, but because fen plants are often phosphorus limited, the plant-fungus interaction could be an important factor in plant competition for phosphorus. In this field study, we determined mycorrhizal colonization rates for 18 fen plant species. Also in the field, we examined the effect of four different forms of phosphorus on the percentage colonization for one fen plant species, Solidago patula. We found that in a species-rich, phosphorus-poor wetland both mycorrhizal and nonmycorrhizal species were common. Nine of ten dicotyledonous species examined formed arbuscular mycorrhizas, while all monocotyledonous species were at most very weakly mycorrhizal. A morphological explanation for this pattern is that the monocots in our study have more extensive aerenchyma, especially in coarse roots. Therefore, monocots are able to transport oxygen to their roots more effectively than dicots. In the organic wetland soil, additional oxygen in the rhizosphere promotes phosphorus mineralization and availability. Two of the monocot species (Typha latifolia and Carex lasiocarpa), which have been described previously as mycorrhizal in other wetland types, are surprisingly nonmycorrhizal in our phosphorus-poor study site, suggesting that a mycorrhizal association would not offer improved phosphorus nutrition to these species. In contrast, our field phosphorus addition decreased mycorrhizal colonization in S. patula, suggesting that one benefit to S. patula of the mycorrhizas is phosphorus uptake.     

土著菌根真菌和混生植物对羊草生长和磷营养的影响

植物种间相互作用直接影响植物生长、根系可塑性及养分吸收,而与植物共生的丛枝菌根真菌可以改变植物个体和种间养分资源的分配,具有协调种间竞争的潜力.以我国北方草甸草原建群种羊草(Leymus chinensis)和混生植物紫花苜蓿(Medicago sativa)及独行菜(Lepidium spetalum)为供试植物,通过模拟盆栽试验,研究了土著菌根真菌和混生植物对羊草生长、根系形态及磷营养的影响.试验结果表明,土著菌根真菌能够与羊草及紫花苜蓿形成良好共生,而独行菜根内基本未形成菌根共生结构.土著菌根真菌显著降低了羊草及独行菜的生物量,但促进了紫花苜蓿的生长;混种紫花苜蓿显著促进了羊草的生长,而混种独行菜则显著抑制了羊草的生长.土著菌根真菌对羊草根系形态的影响表现出与植株生物量类似的趋势,但不同混生植物对羊草根系生长均无显著影响.土著菌根真菌和混生植物对羊草植株磷含量均无显著影响.与混生植物相比,羊草具有较高的比根长和磷吸收能力,这也解释了其负向菌根依赖性.研究证实了菌根真菌和植物种间相互作用均是影响草原优势植物生长和根系发育的重要因素,深入研究其交互作用对于科学管理草地生态系统,维持植物群落的稳定性和生态系统生产力具有重要意义.     

Mycorrhizas and mycorrhizal fungal communities throughout ecosystem development

Background and scope

Plant communities and underlying soils undergo substantial, coordinated shifts throughout ecosystem development. However, shifts in the composition and function of mycorrhizal fungi remain poorly understood, despite their role as a major interface between plants and soil. We synthesise evidence for shifts among mycorrhizal types (i.e., ectomycorrhizas, arbuscular and ericoid mycorrhizas) and in fungal communities within mycorrhizal types along long-term chronosequences that include retrogressive stages. These systems represent strong, predictable patterns of increasing, then declining soil fertility during ecosystem development, and are associated with coordinated changes in plant and fungal functional traits and ecological processes.

Conclusions

Mycorrhizal types do not demonstrate consistent shifts through ecosystem development. Rather, most mycorrhizal types can dominate at any stage of ecosystem development, driven by biogeography (i.e., availability of mycorrhizal host species), plant community assembly, climate and other factors. In contrast to coordinated shifts in soil fertility, plant traits and ecological processes throughout ecosystem development, shifts in fungal communities within and among mycorrhizal types are weak or idiosyncratic. The consequences of these changes in mycorrhizal communities and their function for plant–soil feedbacks or control over long-term nutrient depletion remain poorly understood, but could be resolved through empirical analyses of long-term soil chronosequences.