Belowground Resource Availability Research Articles

Root exudation in a sloping Moso bamboo forest in relation to fine root biomass and traits.

Exudation by fine roots generally varies with their morphological traits, but the effect of belowground resource availability on the root exudation via root morphological traits and biomass remains unknown. We aimed to determine the effects of morphological and physiological traits on root exudation rates and to estimate stand-scale exudation (Estand) by measuring the mass, length, and surface area of fine roots in a Moso bamboo forest. We measured root exudation as well as morphological and physiological traits in upper and lower plots on a slope with different belowground resource availability. The mean (± S.D.) root exudation rates per mass in the upper and lower slope were 0.049 ± 0.047 and 0.040 ± 0.059 mg C g-1 h-1, respectively, which were in the range of exudation found in woody forest ecosystems. We observed significant relationships between root exudation per mass and root respiration, as well as specific root length and surface area. In contrast, exudation per length and area did not correlate with morphological traits. The morphological traits did not differ between slope positions, resulting in no significant difference in root exudation per mass. Fine root biomass, length, and surface area on a unit ground basis were much higher in the lower than those in the upper slope positions. Estand was higher when estimated by mass than by length and area because the morphological effect on exudation was ignored when scaled using mass. Estand was 1.4–2.0-fold higher in the lower than that in upper slope positions, suggesting that the scaling parameters of mass, length, and area determined the Estand estimate more than the exudation rate per mass, length, and area. Regardless of scaling, Estand was much higher in the Moso bamboo forest than in other forest ecosystems because of a large fine-root biomass.

Differential Mechanisms Drive Species Loss Under Artificial Shade and Fertilization in the Alpine Meadow of the Tibetan Plateau.

Fertilization is an effective management strategy to promote community biomass but can simultaneously reduce species diversity in many grassland systems. Shifts in competition for resources have been proposed to explain the decline in plant species diversity due to fertilization, yet the underlying mechanism driving species loss remains controversial. This uncertainty may be driven by variation in aboveground and belowground resource availability. However, experiments simultaneously manipulating both light availability and soil nutrients are rare. Using a 6-year field experiment to manipulate light availability (via shade cloth) and soil nutrients (via fertilizer addition), we tested this resource competition hypothesis in a species-rich alpine meadow by examining the variation of species traits associated with the capacity of light acquisition within these treatments. Our results showed that artificial shade decreased community biomass accumulation whereas fertilization increased it. In contrast, both shade and fertilization reduced species diversity. Extinction of non-Gramineae species (e.g., Fabaceae and Cyperaceae) was the main reason for species diversity decline. Species loss can be explained by the limitation of light availability and predicted by species traits associated with light acquisition capability under fertilization and low light tolerance under artificial shade. Specifically, fertilization eliminated species with lower stature and artificial shade exterminated species with the higher light compensation point (LCP). The findings suggest that light availability is consistently important for plant growth and that low competitiveness for light under fertilization and intolerance of low light conditions under artificial shade trigger species loss process in the alpine meadow. Our experiment helps clarify the mechanisms of how artificial shade and fertilization decreased species diversity and highlight that LCP, which tends to be neglected by most of the studies, is one of the vital drivers in determining species coexistence.

Climatic conditions, not above- and belowground resource availability and uptake capacity, mediate tree diversity effects on productivity and stability

Tree species diversity promotes multiple ecosystem functions and services. However, little is known about how above- and belowground resource availability (light, nutrients, and water) and resource uptake capacity mediate tree species diversity effects on aboveground wood productivity and temporal stability of productivity in European forests and whether the effects differ between humid and arid regions. We used the data from six major European forest types along a latitudinal gradient to address those two questions. We found that neither leaf area index (a proxy for light uptake capacity), nor fine root biomass (a proxy for soil nutrient and water uptake capacity) was related to tree species richness. Leaf area index did, however, enhance productivity, but negatively affected stability. Productivity was further promoted by soil nutrient availability, while stability was enhanced by fine root biomass. We only found a positive effect of tree species richness on productivity in arid regions and a positive effect on stability in humid regions. This indicates a possible disconnection between productivity and stability regarding tree species richness effects. In other words, the mechanisms that drive the positive effects of tree species richness on productivity do not per se benefit stability simultaneously. Our findings therefore suggest that tree species richness effects are largely mediated by differences in climatic conditions rather than by differences in above- and belowground resource availability and uptake capacity at the regional scales.

Fine Root Growth of Black Spruce Trees and Understory Plants in a Permafrost Forest Along a North-Facing Slope in Interior Alaska.

Permafrost forests play an important role in the global carbon budget due to the huge amounts of carbon stored below ground in these ecosystems. Although fine roots are considered to be a major pathway of belowground carbon flux, separate contributions of overstory trees and understory shrubs to fine root dynamics in these forests have not been specifically characterized in relation to permafrost conditions, such as active layer thickness. In this study, we investigated fine root growth and morphology of trees and understory shrubs using ingrowth cores with two types of moss substrates (feather- and Sphagnum mosses) in permafrost black spruce (Picea mariana) stands along a north-facing slope in Interior Alaska, where active layer thickness varied substantially. Aboveground biomass, litterfall production rate, and fine root mass were also examined. Results showed that aboveground biomass, fine root mass, and fine root growth of black spruce trees tended to decrease downslope, whereas those of understory Ericaceae shrubs increased. Belowground allocation (e.g., ratio of fine root growth/leaf litter production) increased downslope in both of black spruce and understory plants. These results suggested that, at a lower slope, belowground resource availability was lower than at upper slope, but higher light availability under open canopy seemed to benefit the growth of the understory shrubs. On the other hand, understory shrubs were more responsive to the moss substrates than black spruce, in which Sphagnum moss substrates increased fine root growth of the shrubs as compared with feather moss substrates, whereas the effect was unclear for black spruce. This is probably due to higher moisture contents in Sphagnum moss substrates, which benefited the growth of small diameter (high specific root length) fine roots of understory shrubs. Hence, the contribution of understory shrubs to fine root growth was greater at lower slope than at upper slope, or in Sphagnum than in feather-moss substrates in our study site. Taken together, our data show that fine roots of Ericaceae shrubs are a key component in belowground carbon flux at permafrost black spruce forests with shallow active layer and/or with Sphagnum dominated forest floor.

Differences in net primary production allocation and nitrogen use efficiency between Moso bamboo and Japanese cedar forests along a slope

ABSTRACT The responses of forest production and nitrogen (N) utilization to belowground resource availability may be stronger in Moso bamboo forests than in woody forests because the former have higher N and water demands than the latter. The aim of this study was to clarify the different effects of mountain slope-induced belowground resource limitation on forest production between a Moso bamboo forest and a Japanese cedar forest. We compared the changes in biomass, net primary production, net primary production allocation, N utilization, N content per organ, and N use efficiency in response to soil resource availability at five different slope positions in each forest type. In the bamboo forest, the aboveground net primary production decreased upslope and the belowground net primary production allocation increased with decreasing upslope soil water availability. In contrast, leaf N content of the bamboo forest increased upslope, and N use efficiency decreased upslope with an increasing upslope soil N mineralization rate. Although a greater N use efficiency has often been reported at higher slope positions due to lower N availability, this phenomenon was observed at lower slope position in the bamboo forest. The soil N supply might exceed the soil N demand at higher slope positions, resulting in lower N use efficiency. The responses observed in the bamboo forest were absent in the Japanese cedar forest. The higher demand for belowground resources and environmental plasticity explained the higher productivity of the bamboo forest.

Site conditions more than species identity drive fine root biomass, morphology and spatial distribution in temperate pure and mixed forests

AimThe establishment of broadleaved - conifer mixtures has had positive effects on productivity. However, it is still not clear the degree to which belowground space occupation contributes to these effects and how root traits of the respective species differ between mixed and pure stands. Thus, the aim of this study was to examine how enrichment of beech forests with conifer species impacts fine root biomass and other root traits across variable site conditions. MethodsWe studied 20 mature pure and mixed stands of European beech (Fagus sylvatica), Norway spruce (Picea abies), and Douglas-fir (Pseudotsuga menziesii), and determined differences in biomass, spatial distribution, and morphology of fine roots (≤2 mm, to 60 cm soil depth) among beech – conifer mixtures and pure stands of the respective species in two regions with differing site conditions in Northern Germany. ResultsSite conditions had strong effects on fine root biomass (FRB) and fine root area index (RAI). Both FRB and RAI were significantly higher across all stand types on the sites with lower belowground resource availability. On the less favorable sites, FRB significantly increased with increasing beech proportion. In contrast to specific root area (SRA) and RAI, specific root length (SRL) was not affected by site conditions but differed significantly among species. No overyielding was observed when FRB was plotted against basal area and/or area potentially available (APA), which reflects the aboveground horizontal space configuration. Spatial fine root biomass patterns of all species in both monospecific and mixed stands changed along the vertical soil profile at both sites. FRB decreased with increasing soil depth, with the exception of beech in mixed stands. ConclusionOur data suggest that both amount and distribution of tree fine roots in pure and mixed stands are highly variable and depend primarily on-site conditions and secondarily on tree species identity. By disentangling soil and species identity effects, we conclude that beech has a remarkable potential for adjusting fine root biomass to cope with unfavorable conditions as compared to spruce and Douglas-fir.

Interactive effects of nitrogen deposition and drought-stress on plant-soil feedbacks of Artemisia californica seedlings

Nitrogen (N) deposition and drought are major drivers of global change that will influence plant-soil feedbacks. We investigated how N availability, N-impacted soil communities and drought affect feedback in seedlings of a drought-deciduous mycorrhizal shrub, Artemisia californica. Seedlings were inoculated with soil from either a high or low deposition site or sterilized inoculum and grown with or without supplemental N and under well-watered or drought-stressed conditions. Inoculum, N and water had interactive effects on feedbacks. Seedlings grown in low deposition inoculum exhibited a neutral to positive feedback under drought and had the highest root to shoot ratios and mycorrhizal colonization. Seedlings inoculated with high N-deposition soil experienced a positive feedback when N fertilized and well-watered, but plants allocated large amounts of biomass to shoots and had a negative response to drought. The soil community mediates plant response to varying belowground resource availability. We found N-impacted communities may reduce mycorrhizal colonization and allocation to roots and provide less protection against drought. Our results highlight the context dependency of plant-soil feedbacks and the potential for climate change and N deposition to have interactive effects on these relationships.

A simple model for the influence of habitat resource availability on lateral clonal spread.

Plant clonal spread is ubiquitous and of great interest, owing both to its key role in plant community assembly and its suitability for plant behaviour research. However, mechanisms that govern spreading distance are not well known. Here we link spacer costs and below-ground competition in a simple model of growth in a homogeneous below-ground environment, in which optimal distance between ramets is based on minimizing the sum of these costs. Using this model, we predict a high prevalence of clonal growth that does not employ spacers in resource-poor environments and a nonlinear increase in spreading distance in response to increasing below-ground resource availability. Analysis of database data on clonal growth in relationship to below-ground resource availability revealed that patterns of the spread based on stolons is compatible with the model's predictions. As expected, model prediction failed for rhizomatous species, where spacer sizes are likely to be selected mainly to play roles other than spread. The model's simplicity makes it useful as a null model in testing hypotheses about the effects of environmental heterogeneity on clonal spread.

Effects of arbuscular mycorrhizal fungi on grassland productivity are altered by future climate and below-ground resource availability

Due to their key position at the soil–root interface, arbuscular mycorrhizal fungi (AMF) may play an important role in determining the impact of global change on plant communities. We studied the effects of mycorrhizal colonization and “future climate” on productivity of model grassland ecosystems. In 2007 and 2008, artificially assembled grassland communities, grown on pasteurized soil inoculated with or free from AMF, were exposed to ambient conditions or to combined elevated atmospheric CO2 concentration and temperature. In the full soil accessibility experiment (“full soil”, 2007) plant roots could explore the total soil volume. In the partial root-exclusion experiment (“partial soil”, 2008), a nylon membrane of 25μm mesh size allowed AMF to explore an additional soil volume that was not accessible to the roots. In the latter experiment, planting density was higher which increased the competition for nutrient uptake by roots compared to the “full soil” experiment.The main findings were the following:•“Future climate” had a positive effect on AMF root colonization.•Under “future climate”, the stimulated AMF suppressed root biomass in both experiments.•AMF increased plant nutrient stocks and above-ground biomass, regardless of the climate, but only in the “partial soil” experiment, when plant competition for nutrients likely was higher and AMF could increase the available nutrient pool by exploring an extra root-free soil compartment.

The effects of belowground resources on aboveground allometric growth in Bornean tree species

Tree height and crown allometries reflect adaptations for resource acquisition and structural stability, as well as plastic responses to a heterogeneous environment. While both light and soil resources limit growth and influence competitive responses in tropical forests, the effects of belowground resources on allometries are less understood, especially within the understory. To characterize outcomes of tree competition along an edaphic resource gradient, we quantified variation in height and crown allometries of six Bornean tree species from contrasting regeneration niches (light-demanding vs. shade-tolerant) on two soil habitats (clay-fine loam and sandy loam) within a 52-ha forest dynamics plot. Using empirically-fit allometric parameters and diameter growth rates from plot census data, we modeled tree height and crown area growth over the projected life span of each species. Based on resource competition theory, we hypothesized that tree species specializing on and populations of generalist species growing on the relatively moister, more fertile clay-fine loam soil habitat would have faster height and crown growth rates, compared to those on the sandy loam habitat, regardless of regeneration niche. Among soil specialists and within generalists of both genera, trees growing on clay-fine loam had taller stems and larger crowns at a given age and faster height and crown area growth rates at most sizes than trees on sandy loam. Differences in height and crown growth were driven by the faster diameter growth rates of trees on clay-fine loam, not by differences in height- and crown-diameter allometries, as trees on sandy loam were significantly taller at a given diameter, and differences in crown allometry were not consistent across soil habitats. Characterizing the height and crown growth responses of trees along resource gradients provides insight into the mechanisms that maintain diversity in tropical forests. Our results point to the importance of adaptive and plastic responses to both above and belowground resource availability in determining the allometric growth of trees and suggest that this diversity of responses may contribute tree species coexistence through competition-based trade-off mechanisms and variation in growth among individuals. Additionally, as the importance estimating natural carbon sequestration increases with the escalating effects of anthropogenic climate change, differences in tree growth and architecture across soil habitats also have implications for the approximation of forest carbon storage on heterogeneous tropical soils.

Giant Sequoia (Sequoiadendron giganteum) Regeneration in Experimental Canopy Gaps

Restoration of giant sequoia populations is a high priority for managers, but few experimental studies have examined the efficacy of restoration treatments. To inform giant sequoia restoration treatment options, we assessed the response of giant sequoia regeneration (germination, mortality, and growth) to experimental gaps within a native giant sequoia grove. We created experimental gaps, ranging in size from 0.05 to 0.4 ha. Following gap creation, we sowed seeds and planted seedlings along north‐south transects across gaps. Transects were planted on paired burned and unburned substrates. The seed‐sowing treatment did not result in a cohort of established seedlings, although the amount of seeds sowed was far short of the potential amount released during intense fires. Mortality of planted seedlings did not vary with gap size (average 25% mortality). However, there was a distinct relationship between gap size and second year seedling growth. The relationship was best modeled with an asymptotic curve for both burned and unburned substrates. Relative seedling growth more than doubled as gap size increased from 0.05 to 0.2 ha, then increases in growth diminished. Growth rates of giant sequoia seedlings saturated above 70% light availability while increasing linearly with belowground resource availability. Long‐lived pioneer species such as giant sequoia require restoration treatments that involve relatively severe disturbances to facilitate cohort establishment and recruitment.

High leaf tissue density grassland species consistently more abundant across topographic and disturbance contrasts in a North American tallgrass prairie

Understanding the coupling between plant functional traits and abundance provides insight into the often hidden forces that structure plant communities. To better understand the coupling between leaf traits and abundance of grassland species in a mesic North American grassland, we measured specific leaf area (SLA) and its two components, tissue density and thickness for 125 grassland species. Plants with high tissue density were more abundant over a 17-year period across a range of environments: uplands, grazed and ungrazed watersheds, and frequently and infrequently burned watersheds. The consistent relationships between leaf tissue density and abundance across ecological contrasts imply that belowground resource availability constrains community composition independent of grazing and burning regimes. Leaf tissue density did not explain species abundance in lowlands, where belowground resources are the highest. Neither did it explain the differential abundance of species between grazing or fire frequency contrasts, suggesting that changes in burning or grazing select for species based on other traits. Relative to leaf tissue density, SLA was a poor predictor of abundance, reinforcing a long-observed—but often ignored—call that measurements of SLA need to be coupled with thickness measurements in order to effectively predict the performance of species. More generally, future research needs to investigate which belowground resources control community composition in the grassland and whether the importance of water or nutrients change with burning and grazing.

Variation in leaf stomatal traits of 28 tree species in relation to gas exchange along an edaphic gradient in a Bornean rain forest

Quantifying variation in functional traits associated with shifts in the species composition of plant communities along resource gradients helps identify environmental attributes important for community assembly. Stomates regulate the balance between carbon assimilation and water status in plants. If environmental attributes affecting photosynthetic water-use efficiency govern species distribution along an edaphic gradient, then adaptive variation in stomatal traits of plant species specializing on different soils should reflect belowground resource availability. • We tested this hypothesis by quantifying stomatal trait variation in understory saplings of 28 Bornean tree species in relation to gas exchange and water-use efficiency (WUE). • Comparisons between congeneric specialists of the more fertile, moister clay and the less fertile, well-drained sandy loam revealed little evidence of similar shifts in stomatal traits across genera, nor was stomatal pore index correlated with g(max), A(max), or WUE (A(max)/g(max) or Δ(13)C), suggesting that stomates may be overbuilt in these shaded juveniles. A(max) was higher on sandy loam, likely due to higher understory irradiance there, but there were no other significant differences in gas exchange or WUE. • Despite substantial diversity in stomatal anatomy, there were few strong relationships between stomatal, photosynthetic, and WUE traits in relation to soil resources. Routine differences in water availability therefore may not exert a dominant control on the distributions of these Bornean tree species. Furthermore, the clades represented by these 12 genera may possess alternative functional designs enabling photosynthetic WUE that is sufficient to these humid, understory environments, due to whole plant-functional integration of stomatal traits with other, unmeasured traits influencing gas exchange.

Performance Trade‐offs Driven by Morphological Plasticity Contribute to Habitat Specialization of Bornean Tree Species

ABSTRACTGrowth‐survival trade‐offs play an important role in niche differentiation of tropical tree species in relation to light‐gradient partitioning. However, the mechanisms that determine differential species performance in response to light and soil resource availability are poorly understood. To examine responses to light and soil nutrient availability, we grew seedlings of five tropical tree species for 12 mo at < 2 and 18 percent full sunlight and in two soil types representing natural contrasts in nutrient availability within a lowland dipterocarp forest in North Borneo. We chose two specialists of nutrient‐rich and nutrient‐poor soils, respectively, and one habitat generalist. Across all species, growth was higher in high than low light and on more nutrient rich soil. Although species differed in growth rates, the ranking of species, in terms of growth, was consistent across the four treatments. Nutrient‐rich soils improved seedling survival and increased growth of three species even under low light. Slower‐growing species increased root allocation and reduced specific leaf area (SLA) and leaf area ratio (LAR) in response to decreased nutrient supply. All species increased LAR in response to low light. Maximum growth rates were negatively correlated with survival in the most resource‐limited environment. Nutrient‐poor soil specialists had low maximum growth rates but high survival at low resource availability. Specialists of nutrient‐rich soils, plus the habitat generalist, had the opposite suite of traits. Fitness component trade‐offs may be driven by both light and belowground resource availability. These trade‐offs contribute to differentiation of tropical tree species among habitats defined by edaphic variation.

Effects of above- and belowground partial harvest disturbance on growth and water status of residual sugar maple

Partial forest harvesting is known to modify both above- and belowground resource availability and may result in direct and indirect stress to the residual trees as a result of machinery traffic and sudden changes in irradiance. We studied sugar maple (Acer saccharum Marsh.) trees in stands that had undergone a selection harvest 11 years before sampling to verify whether sudden increases in light availability and soil disturbance caused by machinery influence growth rates and lead to water stress. We selected trees that had experienced either no disturbance from partial harvest, soil disturbance only, sudden increases in light availability only or both disturbances. We analyzed stem radial growth rates and stable carbon isotope composition (delta(13)C) of stem wood with an annual resolution from 10 years before partial harvest until 10 years after partial harvest. Disturbances from partial harvest did not negatively affect growth rates or tree water status. Although trees that experienced increased light availability had higher (less negative) delta(13)C after harvest (indicating increased water-use efficiency), they also had higher growth rates, suggesting that they experienced no pronounced water stress. Trees subjected to soil disturbance showed no sign of water stress. These results may partly be associated with favorable growth conditions (abundant precipitation and mild temperature) in the years following harvest and could differ from results that would be observed under more severe climatic conditions.

Ontogenetic variation in the relative influence of light and belowground resources on European beech seedling growth

We used height growth data from a 7-year field experiment with European beech (Fagus sylvatica L.) seedlings to test the hypothesis that the effects of above- and belowground resources on height growth depend on seedling size and age. Photosynthetically active radiation (PAR) was determined by hemispheric photography, and estimates of fine root biomass of the overstory trees were used as an inverse proportional surrogate for belowground resource availability. For recently germinated seedlings growing under the canopy of Picea abies (L.) Karst., belowground resource availability affected height growth more than light. During subsequent stages of seedling development, apart from initial seedling size, PAR increasingly determined seedling growth. Besides initial size, seedling age determined the effects of above- and belowground resources on seedling height growth. In seedlings identical in initial size but differing in age, the increase in height growth with increasing PAR was greater in older seedlings than in younger seedlings. The ranking of seedling height by year showed that small differences in size at the end of the first growing season resulted in continuously increasing differences during the following years. Mortality data indicated that the chances of a seedling surviving intraspecific competition was strongly determined by its dominance ranking within the first 5 years after establishment.

Interspecific demographic trade‐offs and soil‐related habitat associations of tree species along resource gradients

1 Interspecific relationships between fundamental demographic rates, often called demographic trade-offs, emerge from constraints within individuals related to morphology, physiology and resource allocation. Plant species that grow fast in high light usually have high mortality in shade, and this well-established relationship in part defines a species’ successional niche. More generally, this relationship represents a trade-off between a species’ ability to grow quickly to exploit abundant resources vs. avoiding mortality when resources are less plentiful, but few studies have described this demographic trade-off with respect to environmental factors other than light. 2 Using demographic data from 960 tree species in Bornean rain forest, we examined the evidence for an interspecific demographic trade-off between fast growth and low mortality and its variation among habitats defined by variation in soil fertility and moisture. Such a trade-off could contribute to sorting of tree species among habitats and partly explain the striking patterns of species’ edaphic associations in this and other forests. 3 We found strong evidence for this demographic trade-off, both within the same habitat and when growth on edaphically rich habitats was compared with mortality on a habitat with lower below-ground resource availability. 4 The slope of the growth-mortality relationship varied among habitats, being steepest on the habitat lowest in below-ground resources. For species with the fastest potential growth rates, mortality was higher on this habitat than at comparable growth rates on the three more edaphically rich habitats, providing a possible mechanism by which fast-growing species may be eliminated from the poorest habitat. Adaptations for fast growth may entail a greater mortality risk, if inherently fast-growing species fail to maintain a positive C-balance when below-ground resources are scarce. 5 Conversely, for species with the slowest potential growth rates, the highest species’ mortality rates occurred on the habitats with greatest below-ground resource availability, implying that slow-growing species may have a competitive disadvantage in resource-rich environments. 6 Synthesis. Differences among habitats in the steepness of this trade-off may sort species into different habitats along this edaphic gradient, whereas on the same soil, this demographic trade-off could facilitate coexistence of at least some species in this forest. Thus, by generating emergent demographic trade-offs that vary along resource gradients, plant life-history strategies can influence species diversity and distribution.

Soil nutrients influence spatial distributions of tropical tree species.

The importance of niche vs. neutral assembly mechanisms in structuring tropical tree communities remains an important unsettled question in community ecology [Bell G (2005) Ecology 86:1757-1770]. There is ample evidence that species distributions are determined by soils and habitat factors at landscape (<10(4) km(2)) and regional scales. At local scales (<1 km(2)), however, habitat factors and species distributions show comparable spatial aggregation, making it difficult to disentangle the importance of niche and dispersal processes. In this article, we test soil resource-based niche assembly at a local scale, using species and soil nutrient distributions obtained at high spatial resolution in three diverse neotropical forest plots in Colombia (La Planada), Ecuador (Yasuni), and Panama (Barro Colorado Island). Using spatial distribution maps of >0.5 million individual trees of 1,400 species and 10 essential plant nutrients, we used Monte Carlo simulations of species distributions to test plant-soil associations against null expectations based on dispersal assembly. We found that the spatial distributions of 36-51% of tree species at these sites show strong associations to soil nutrient distributions. Neutral dispersal assembly cannot account for these plant-soil associations or the observed niche breadths of these species. These results indicate that belowground resource availability plays an important role in the assembly of tropical tree communities at local scales and provide the basis for future investigations on the mechanisms of resource competition among tropical tree species.

Compensation among root classes in Phaseolus vulgaris L.

The activity of soil pathogens, competition for assimilates, and the changing availability of below-ground resources make root systems subject to a continuous and dynamic process of formation and loss of both fine and coarse roots. As hypocotyl borne roots appear later than other root classes, they may serve to functionally replace basal and primary roots lost to biotic and abiotic stress. Using common bean (Phaseolus vulgaris L.), we conducted experiments in solution and solid media culture with treatments involving the removal of part of the root system (basal, hypocotyl borne or primary roots), phosphorus availability, and depth of seeding to test the hypothesis that there are compensation mechanisms among basal, hypocotyl borne and primary roots to cope with the loss of part of the root system. The root system was highly plastic in response to root excision, which resulted in the maintenance of below-ground biomass accumulation. In most cases, this compensation among root classes was enough to maintain plant performance in both phosphorus sufficient and phosphorus stressed plants. Removal of a specific root class induced an increase in the growth of the remaining root classes. All root classes, but especially the primary root, contributed to the compensation mechanism in some way. Primary roots represented around 10% of the root system in control plants and this proportion increased dramatically (up to 50%) when other root classes were removed. In contrast, negligible compensatory re-growth was observed following removal of the primary root. Greater planting depth increased the production of hypocotyl borne roots at the expense of basal roots. The proportion of hypocotyl borne roots increased from 25% of the whole root system when seeds were placed at a depth of 2 cm to 30% when they were placed at 5 cm and to 38% when placed at 8 cm, with corresponding decreases in the proportion represented by basal roots. The common feature of our observations is the innate ability of the root system for its own regeneration. Total root biomass maintained strict allometric relationships with total shoot biomass in all treatments. Re-stabilization of root to shoot balance after partial root loss is governed by overall plant size following allometric relationships similar to undisturbed plants. However, the pattern of this root regeneration was not uniform since the way the three root classes compensated each other after the removal of any one of them varied among the different growth media and phosphorus supply conditions. The resulting changes in root architecture could have functional significance for soil resource acquisition.

Growth limitations of planted conifers regenerating under Montane Alternative Silviculture Systems (MASS): Seven-year results

As part of the Montane Alternative Silviculture Systems (MASS) project, this study investigates limits on the growth of montane conifers resulting from varying overstory retention under conventional and alternative silvicultural systems. After harvesting treatments were complete in 1993, Abies amabilis (amabilis fir) and Tsuga heterophylla (western hemlock) seedlings were spring planted in replicated blocks of shelterwood (SW), patch cut (PC), green tree retention (GT) and clearcut (CC) systems. In addition, sub-plots were established within each silvicultural system in which fertilization (at planting) and vegetation control post-planting treatments were applied alone and in combination to test the extent to which growth limitations are related to nutrient availability and vegetative competition. The impact of overstory retention was most pronounced in the reduced light environment of the SW where height growth after seven years was 26-30% lower in both species compared to the untreated CC, GT and PC systems. Although the effect on growth of both species in the SW was mitigated somewhat by fertilization and vegetation control treatments, amabilis fir did not attain free-to-grow height (1.3 m) regardless of post-planting treatment. Time to free-to-grow height in the more open silvicultural systems was reduced in both fir and hemlock with fertilization and vegetation control alone and in combination, except in the CC where the initial growth response to fertilization alone was diminished by the end of the seven-year study. In contrast to fertilization, the effects of vegetation control on height growth were not apparent until three to five years and seven years after planting in the CC and GT, PC, SW, respectively. Combining vegetation control and fertilization had an additive effect on growth in amabilis fir but not in western hemlock. The effect of silvicultural systems and post-planting treatments on the two species illustrate that both above- and below-ground resource availability (light and nutrients) availability was potentially limiting to growth, particularly in the shelterwood treatment. Key words: MASS, silviculture systems, regeneration, Tsuga heterophylla, Abies amabilis, shelterwood, patch cut, green tree reten-tion, clearcut, fertilizer, vegetation control