Longevity and turnover of roots in the shortgrass steppe: influence of diameter and depth

We used minirhizotrons to determine patterns of root longevity andturnover for the perennial bunchgrass Bouteloua gracilisinthe shortgrass steppe of eastern Colorado, USA. We hypothesized that rootlongevity would be partially controlled by root diameter, following previouslyobserved patterns in woody plants. In addition, we hypothesized that rootturnover would be greatest in surface soil horizons and decrease with depth dueto variation in soil moisture availability and temperature. Root longevity wascorrelated with root diameter. Median life span of roots > 0.4mm was approximately 320 days, while roots < 0.2mmhad a median life span of 180 days. There was approximately a 6%decreasein the likelihood of mortality with a 0.10-mm increase inroot diameter, controlling for the effect of depth in the soil profile. Rootlength production and mortality were highest in the upper20 cm of the soil profile and decreased with depth.However,because root length density also decreased with depth, there were nosignificantdifferences in turnover rate of root length among sampling intervals. Turnoverwas approximately 0.86 yr^–1 based on root length production,while turnover was 0.35 yr^–1 using root length mortality as ameasurement of flux. The imbalance between turnover estimates may be aconsequence of the time the minirhizotrons were in place prior to imaging or mayresult from our lack of over-winter measures of mortality. Our worksuggests that Bouteloua gracilis roots have complex lifehistory strategies, similar to woody species. Some portion of the root systemishighly ephemeral, while slightly larger roots persist much longer. Thesedifferences have implications for belowground carbon and nitrogen cycles in theshortgrass steppe.     

Waterlogging responses in dune,swale and marsh populations of Spartina patens under field conditions

Summary Soil waterlogging responses were examined in three Spartina patens populations along a steep flooding gradient in coastal Louisiana. Root anatomy and physiological indicators of anaerobic metabolism were examined to identify and compare flooding responses in dune, swale and marsh populations, while soil physicochemical factors were measured to characterize the three habitats. Soil waterlogging increased along the gradient from dune to marsh habitats and was accompanied by increases in root porosity (aerenchyma). Aerenchyma in marsh roots was apparently insufficient to provide enough oxygen for aerobic respiratory demand, as indicated by high root alcohol dehydrogenase activities and low energy charge ratios. Patterns of root metabolic indicators suggest that dune and swale roots generally respired aerobically, while anaerobic metabolism was important in marsh roots. However, in each population, relatively greater soil waterloging was accompanied by differences in enzyme activities leading to malate accumulation. In dune and swale roots under these circumstances, depressed adenylate energy charge ratios may have been the result of an absence of increased ethanol fermentation. These trends suggest that: 1) Aerenchyma formation was an important, albeit incomplete, long-term adaptation to the prevalent degree of soil waterlogging. 2) All populations adjusted root metabolism in response to a relative (short-term) increase in soil waterlogging.     

The effect of nitrogen fertilization on the phenology of roots in a barrier island sand dune community

Little work has been done on the phenology of root growth and senescence largely due to methodological difficulties. The application of minirhizotron technology has enabled the tracking of individual roots through an entire growing season. As a result, direct measures of mortality, root growth, and an analysis of cohorts can be obtained. This study examined the belowground response of vegetation in a nutrient limited system to nitrogen addition. Small plots on a 36 year old dune on Hog Island, a barrier island in the Virginia Coast Reserve Long Term Ecological Research Site, were fertilized with nitrogen. Minirhizotron tubes were installed in each fertilized and control plot. Each tube was sampled monthly for nine months, March through October of 1992. Root length density increased throughout the growing season with the greatest root length density in the top 20 cm of the soil profile. The fertilized plots had greater root length densities (14.1 mm cm^-2) than the unfertilized plots (2.9 mm cm^-2). There was no significant depth × treatment interaction. Root mortality did not significantly change with fertilization. The largest loss of roots for a cohort occurred within the first month. The dune grassland community did not respond to fertilization with large changes in root distribution or increases in mortality in this study.     

Annual soil respiration in broadleaf forests of northern Wisconsin: influence of moisture and site biological,chemical, and physical characteristics

Soil temperature and moisture influence soil respiration at a range of temporal and spatial scales. Although soil temperature and moisture may be seasonally correlated, intra and inter-annual variations in soil moisture do occur. There are few direct observations of the influence of local variation in species composition or other stand/site characteristics on seasonal and annual variations in soil moisture, and on cumulative annual soil carbon release. Soil climate and soil respiration from twelve sites in five different forest types were monitored over a 2-year period (1998–1999). Also measured were stand age, species composition, basal area, litter inputs, total above-ground wood production, leaf area index, forest floor mass, coarse and fine root mass, forest floor carbon and nitrogen concentration, root carbon and nitrogen concentration, soil carbon and nitrogen concentration, coarse fraction mass and volume, and soil texture. General soil respiration models were developed using soil temperature, daily soil moisture, and various site/soil characteristics. Of the site/soil characteristics, above-ground production, soil texture, roots + forest floor mass, roots + forest floor carbon:nitrogen, and soil carbon:nitrogen were significant predictors of soil respiration when used alone in respiration models; all of these site variables were weakly to moderately correlated with mean site soil moisture. Daily soil climate data were used to estimate the annual release of carbon (C) from soil respiration for the period 1998–1999. Mean annual soil temperature did not differ between the 2 years but mean annual soil moisture was approximately 9% lower in 1998 due to a summer drought. Soil C respired during 1998 ranged from 8.57 to 11.43 Mg C ha⁻¹ yr⁻¹ while the same sites released 10.13 and 13.57 Mg C ha⁻¹ yr⁻¹ in 1999; inter-annual differences of 15.41 and 15.73%, respectively. Among the 12 sites studied, we calculated that the depression of soil respiration linked to the drought caused annual differences of soil respiration from 11.00 to 15.78%. Annual estimates of respired soil C decreased with increasing site mean soil moisture. Similarly, the difference of respired carbon between the drought and the non-drought years generally decreased with increasing site mean soil moisture.     

ROOT PRODUCTION IN FOUR COMMUNITIES IN THE GREAT DISMAL SWAMP

A sequential coring approach was used to measure root biomass and production over 1 year in four different communities within the Great Dismal Swamp. A second method, an implanted bag technique, was also used to measure root production, and values were generally lower using this technique. On all sites, fine roots were the most dynamic root component. Both biomass (1,887 g/m²) and production (354–989 g m ² yr^-1) were highest on the mixed hardwood site, the least flooded site, and second highest on the cedar site, the site with the longest duration of soil saturation (1,033 g/m² and 274–366 g m^-2 yr^-1). The maple-gum (696 g/m² and 59–91 g m^-2 yr^-1) and cypress (824 g/m² and 68–308 g m^-2 yr^-1) sites had similarly low amounts of biomass and rates of production. Environmental parameters that influenced production include frequency and duration of flooding, and soil type. Peaks in belowground production were observed on the most productive sites (mixed hardwood and cedar) in summer and late fall-winter; the other two sites exhibited little seasonal variability. The least flooded stand appears to allocate a greater percentage of net primary production belowground than the more extensively flooded stands. The ratio of above- and belowground allocation appears to change in response to a flooding gradient. This has major implications for ecosystem functions as carbon allocation patterns determine the array of litter types generated (leaves vs. roots) which affect decomposition rates and nutrient availability.     

ORGANIC MATTER DYNAMICS IN FOUR SEASONALLY FLOODED FOREST COMMUNITIES OF THE DISMAL SWAMP

Budgets of organic matter dynamics for plant communities of the Great Dismal Swamp were developed to summarize an extensive data base, determine patterns of biomass allocation, transfer and accumulation, and make comparisons with other forested wetlands. Aboveground net primary production on the flooded sites (1,050–1,176 g m^-2 yr^-1) was significantly greater than on a rarely flooded site (831 g m^-2 yr^-1). Estimates of belowground net primary production were comparable to aboveground production on flooded sites (824–1,221 gm^-2 yr^-1). However, productivity was nearly three times greater belowground than aboveground on the rarely flooded site (2,256 g m^-2 yr^-1). Aboveground productivity in Dismal Swamp forests is relatively high compared to other forested wetlands. This is attributed to the timing and periodic nature of flood events. Fine root turnover is shown to be an important source of soil organic matter. Estimates indicate that roots contribute about 60% of the annual increment to soil organic matter. Leaflitter contributes 6–28% and wood debris contributes 5–15%. Comparisons with other forested wetlands suggest that detritus accounts for greater than half of the total organic matter (living + dead) in many wetland systems.     

Estimation of carbon allocation to the roots from soil respiration measurements of oil palm

CO₂ flux from the soil was measured in situ under oil palms in southern Benin. The experimental design took into account the spatial variability of the root density, the organic matter in the soil-palm agrosystem and the effect of factors such as the soil temperature and moisture.Measurements of CO₂ release in situ, and a comparison with the results obtained in the laboratory from the same soil free of roots, provided an estimation of the roots contribution to the total CO₂ flux. The instantaneous values for total release in situ were between 3.2 and 10.0 mol CO₂ m^-2 s^-1. For frond pile zones rich in organic matter, and around oil palm trunks, root respiration accounted for 30% of the efflux when the soil was at field capacity and 80% when the soil was dry with a pF close to 4.2. This proportion remained constant in interrow zones at around 75%, irrespective of soil moisture.Subsequently carbon allocation to the roots was determined. Total CO₂ release over a year was 57 Mg of CO₂ ha^-1 yr^-1 (around 1610 g of C per m² per year), and carbon allocation to the roots was approximately 53 Mg of CO₂ ha^-1 yr^-1 of which approximately 13 Mg CO₂ ha^-1 yr^-1 (25%) was devoted to turn-over and 40 Mg CO₂ ha^-1 yr^-1 (75%) to respiration.     

Nitrogen Controls on Fine Root Substrate Quality in Temperate Forest Ecosystems

Nitrogen controls on fine root substrate quality (that is, nitrogen and carbon-fraction concentrations) were assessed using nitrogen availability gradients in the Harvard Forest chronic nitrogen addition plots, University of Wisconsin Arboretum, Blackhawk Island, Wisconsin, and New England spruce-fir transect. The 27 study sites encompassed within these four areas collectively represented a wide range of nitrogen availability (both quantity and form), soil types, species composition, aboveground net primary production, and climatic regimes. Changes in fine root substrate quality among sites were most frequently and strongly correlated with nitrate availability. For the combined data set, fine root nitrogen concentration increased (adjusted R ² = 0.46, P < 0.0001) with increasing site nitrate availability. Fine root “extractive” carbon-fraction concentrations decreased (adjusted R ² = 0.32, P < 0.0002), “acid-soluble” compounds increased (adjusted R ² = 0.35, P < 0.0001), and the “acid-insoluble” carbon fraction remained relatively high and stable (combined mean of 48.7 ± 3.1% for all sites) with increasing nitrate availability. Consequently, the ratio of acid-insoluble C–total N decreased (adjusted R ² = 0.40, P < 0.0001) along gradients of increasing nitrate availability. The coefficients of determination for significant linear regressions between site nitrate availability and fine root nitrogen and carbon-fraction concentrations were generally higher for sites within each of the four study areas. Within individual study sites, tissue substrate quality varied between roots in different soil horizons and between roots of different size classes. However, the temporal variation of fine root substrate quality indices within specific horizons was relatively low. The results of this study indicate that fine root substrate quality increases with increasing nitrogen availability and thus supports the substrate quality component of a hypothesized conceptual model of nitrogen controls on fine root dynamics that maintains that fine root production, mortality, substrate quality, and decomposition increase with nitrogen availability in forest ecosystems in a manner that is analogous to foliage.     

Net primary production and net ecosystem production of a boreal black spruce wildfire chronosequence

Net primary production (NPP) was measured in seven black spruce (Picea mariana (Mill.) BSP)‐dominated sites comprising a boreal forest chronosequence near Thompson, Man., Canada. The sites burned between 1998 and 1850, and each contained separate well‐ and poorly drained stands. All components of NPP were measured, most for 3 consecutive years. Total NPP was low (50–100 g C m^?2 yr^?1) immediately after fire, highest 12–20 years after fire (332 and 521 g C m^?2 yr^?1 in the dry and wet stands, respectively) but 50% lower than this in the oldest stands. Tree NPP was highest 37 years after fire but 16–39% lower in older stands, and was dominated by deciduous seedlings in the young stands and by black spruce trees (>85%) in the older stands. The chronosequence was unreplicated but these results were consistent with 14 secondary sites sampled across the landscape. Bryophytes comprised a large percentage of aboveground NPP in the poorly drained stands, while belowground NPP was 0–40% of total NPP. Interannual NPP variability was greater in the youngest stands, the poorly drained stands, and for understory and detritus production. Net ecosystem production (NEP), calculated using heterotrophic soil and woody debris respiration data from previous studies in this chronosequence, implied that the youngest stands were moderate C sources (roughly, 100 g C m^?2 yr^?1), the middle‐aged stands relatively strong sinks (100–300 g C m^?2 yr^?1), and the oldest stands about neutral with respect to the atmosphere. The ecosystem approach employed in this study provided realistic estimates of chronosequence NPP and NEP, demonstrated the profound impact of wildfire on forest–atmosphere C exchange, and emphasized the need to account for soil drainage, bryophyte production, and species succession when modeling boreal forest C fluxes.     

Effects of soil phosphorus availability,temperature and moisture on soil respiration in Eucalyptus pauciflora forest

Rates of soil respiration (CO₂ efflux) were measured for a year in a mature Eucalyptus pauciflora forest in unfertilized and phosphorus-fertilized plots. Soil CO₂ efflux showed a distinct seasonal trend, and average daily rates ranged from 124 to 574 mg CO₂ m^–2 hr^–1. Temperature and moisture are the main variables that cause variation in soil CO₂ efflux; hence their effects were investigated over a year so as to then differentiate the treatment effect of phosphorus (P) nutrition.Soil temperature had the greatest effect on CO₂ efflux and exhibited a highly significant logarithmic relationship (r² = 0.81). Periods of low soil and litter moisture occurred during summer when temperatures were greater than 10 °C, and this resulted in depression of soil CO₂ efflux. During winter, when temperatures were less than 10 °C, soil and litter moisture were consistently high and thus their variation had little effect on soil CO₂ efflux. A multiple regression model including soil temperature, and soil and litter moisture accounted for 97% of the variance in rates of CO₂ efflux, and thus can be used to predict soil CO₂ efflux at this site with high accuracy. Total annual efflux of carbon from soil was estimated to be 7.11 t C ha^–1 yr^–1. The model was used to predict changes in this annual flux if temperature and moisture conditions were altered. The extent to which coefficients of the model differ among sites and forest types requires testing.Increased soil P availability resulted in a large increase in stem growth of trees but a reduction in the rate of soil CO₂ efflux by approximately 8%. This reduction is suggested to be due to lower root activity resulting from reduced allocation of assimilate belowground. Root activity changed when P was added to microsites within plots, and via the whole tree root system at the plot level. These relationships of belowground carbon fluxes with temperature, moisture and nutrient availability provide essential information for understanding and predicting potential changes in forest ecosystems in response to land use management or climate change.     

Translocation,remineralization, and turnover of nitrogen in the roots and rhizomes of Spartina alterniflora (Gramineae)

We used ¹⁵N to quantify rates of N translocation from aerial to belowground tissues, foliar leaching, and turnover and production of root and rhizome biomass in the plant-sediment system of short Spartina alterniflora areas of Great Sippewissett Marsh, Massachusetts. Decay of belowground tissues in litterbag incubations at 1- and 10-cm depths resulted in 80% remineralization of the original plant (¹⁵N-labeled) N and 20% burial after 3 years. Translocation of ¹⁵N from plant shoots in hydrologically controlled laboratory lysimeters maintained under field conditions was 38% of the aboveground pool while leaching of N was 10% from June to October. Most of the translocated N was not retranslocated to new aboveground growth in December but appeared to be either remineralized or buried in the sediment. Injection of ¹⁵N into field stands of grass showed initially high incorporation into plants followed by a continuous decline over the next 7 years yielding a gross tumover time of 1.5–1.6yr. Correcting the gross N turnover for recycling of label via translocation and uptake of remineralized label during this period, a net root and rhizome turnover time of 1.0–1.1 yr was obtained. Combining the turnover time with independent estimates of seasonal belowground biomass yielded an estimate of belowground production of 929–1,022 g C m⁻² yr⁻¹, similar to measurements by traditional biomass harvest, CO² based budgets and models for comparable areas of this marsh. Integration of the production and nitrogen balance estimates for short Spartina marsh yielded translocation, 1.4 g N m⁻² yr⁻¹, leaching, 0.4 g N m⁻² yr⁻¹, remineralization, 14.9–16.3 g N m⁻² yr⁻¹, and burial, 3.7–4.1 g N m⁻² yr⁻¹.     

Effects of fire,mowing and nitrogen addition on root characteristics in tall-grass prairie

Abstract. Root harvests and root windows were used to study the influence of fire, mowing and nitrogen additions on root lengths, biomass, and nitrogen content in tall-grass prairie. Four years of nitrogen additions (10 g m² yr^?1) increased below-ground mass by 15 % and nitrogen concentration in that mass by 77 %. In general, live roots and rhizomes exhibited greater increases in nitrogen concentrations than detrital roots and rhizomes. After four years of treatment, live roots and rhizomes immobilized an additional 1.5 to 5 g/m² of nitrogen, depending upon specific treatment, while dead roots and rhizomes immobilized an additional 3 to 3.5 g/m². Average root growth parameters, as measured with root windows, were positively correlated with above-ground peak foliage biomass; however, the only significant correlation was between average new root growth and above-ground peak foliage biomass (r = 0.73, p ≤ 0.04). Root growth and decay, as measured by annual mean values for eight root windows over a four year interval, were insensitive to climatic and treatment effects.     

Are patterns in nutrient limitation belowground consistent with those aboveground: results from a 4 million year chronosequence

Accurately predicting the effects of global change on net carbon (C) exchange between terrestrial ecosystems and the atmosphere requires a more complete understanding of how nutrient availability regulates both plant growth and heterotrophic soil respiration. Models of soil development suggest that the nature of nutrient limitation changes over the course of ecosystem development, transitioning from nitrogen (N) limitation in ‘young’ sites to phosphorus (P) limitation in ‘old’ sites. However, previous research has focused primarily on plant responses to added nutrients, and the applicability of nutrient limitation-soil development models to belowground processes has not been thoroughly investigated. Here, we assessed the effects of nutrients on soil C cycling in three different forests that occupy a 4 million year substrate age chronosequence where tree growth is N limited at the youngest site, co-limited by N and P at the intermediate-aged site, and P limited at the oldest site. Our goal was to use short-term laboratory soil C manipulations (using ¹⁴C-labeled substrates) and longer-term intact soil core incubations to compare belowground responses to fertilization with aboveground patterns. When nutrients were applied with labile C (sucrose), patterns of microbial nutrient limitation were similar to plant patterns: microbial activity was limited more by N than by P in the young site, and P was more limiting than N in the old site. However, in the absence of C additions, increased respiration of native soil organic matter only occurred with simultaneous additions of N and P. Taken together, these data suggest that altered nutrient inputs into ecosystems could have dissimilar effects on C cycling above- and belowground, that nutrients may differentially affect of the fate of different soil C pools, and that future changes to the net C balance of terrestrial ecosystems will be partially regulated by soil nutrient status.     

Belowground nitrogen dynamics in relation to hurricane damage along a tropical dry forest chronosequence

Understanding and predicting the responses of plant communities to multiple overlapping disturbances remains a challenging task. Hurricane Wilma represents a large, yet infrequent type of disturbance that was superimposed on an existing disturbance gradient of time since fire. We examined disturbance and recovery patterns in response to these overlapping disturbances by measuring how canopy structure, fine roots, mycorrhizae, and soil nitrogen dynamics, varied along a fire chronosequence in the 2 years after Hurricane Wilma. Hurricane damage increased canopy openness in all seral stages. In the early-seral stage, canopy openness returned to pre-hurricane conditions within 2 years, whereas canopy openness in the late-seral stage remained significantly higher throughout the study. We observed no significant change in root length density in the early- and mid-seral stages. However, in the late-seral stage, root length density was significantly reduced immediately after the hurricane and remained so 2 years after the hurricane. In the late-seral stage, we also observed a significant reduction in percent soil nitrogen and a significant increase in soil nitrogen isotopic composition (δ¹⁵N) values, indicating a loss of soil nitrogen. In contrast, in the early- and mid-seral stages, there were no significant changes in percent nitrogen or soil δ¹⁵N values. Results from this study suggest that forest fire disturbance history influences responses to hurricane damage. Moreover, feedbacks between aboveground and belowground processes have the potential to influence forest recovery.     

The dependence of root system properties on root system biomass of 10 North American grassland species

Dependence of the properties of root systems on the size of the root system may alter conclusions about differences in plant growth in different environments and among species. To determine whether important root system properties changed as root systems aged and accumulated biomass, we measured three important properties of fine roots (tissue density, diameter, and C:N) and three biomass ratios (root:shoot, fine:coarse, and shallow:deep) of monocultures of 10 North American grassland species five times during their second and third years of growth. With increasing belowground biomass, root tissue density increased and diameter decreased. This may reflect cortical loss associated with the aging of roots. For non-legumes, fine root C:N decreased with increasing root biomass, associated with decreases in soil solution NO^{3 –} concentrations. No changes in fine root C:N were detected with increasing belowground biomass for the two legumes we studied. Among all 10 species, there were generally no changes in the relative amounts of biomass in coarse and fine roots, root:shoot, or the depth placement of fine roots in the soil profile as belowground biomass increased. Though further research is needed to separate the influence of root system size, age of the roots, and changes in nutrient availability, these factors will need to be considered when comparing root functional traits among species and treatments.     

Nitrogen cycle of tropical perennial crops under shade trees

The distribution and fluxes of nitrogen in some parts of a coffee plantation under shade were studied at a typical mountain (1380 m a sl) location in Venezuela. The amounts of nitrogen in the soil to 60 cm depth are by far the largest nitrogen store, reaching a total of 49 000 kg ha^?1. The nitrogen flow associated with litterfall was dominated by the shade-tree fraction accounting for a transfer of 86 kg ha^?1 yr^?1 of the total 189 kg ha^?1 yr^?1. The rapid decomposition of this litter, although showing a phase of nitrogen accumulation, is an important source of nitrogen to the roots of coffee which occupy preferentially the upper 30 cm of soil and even the litter layer itself. Some evidence of synchrony was found between the peaks of nitrogen transfer to the soil by litter and the periods of high nitrogen demand by the crop plants. It is proposed that the system can amply compensate the nitrogen outputs by harvest (17 kg ha^?1 yr^?1) with a subsidy from the shade trees.Erythrina sp. andInga sp. are potential nitrogen fixers although we found no active sites during the dry period sampled. The average litter decomposition constant, k, expressed in terms of nitrogen, was estimated as 4.5, equivalent to a half-life of approximately two months.     

Do increased summer precipitation and N deposition alter fine root dynamics in a Mojave Desert ecosystem?

Climate change is expected to impact the amount and distribution of precipitation in the arid southwestern United States. In addition, nitrogen (N) deposition is increasing in these regions due to increased urbanization. Responses of belowground plant activity to increases in soil water content and N have shown inconsistent patterns between biomes. In arid lands, plant productivity is limited by water and N availability so it is expected that changes in these factors will affect fine root dynamics. The objectives of this study were to quantify the effects of increased summer precipitation and N deposition on fine root dynamics in a Mojave Desert ecosystem during a 2‐year field experiment using minirhizotron measurements. Root length density, production, and mortality were measured in field plots in the Mojave Desert receiving three 25 mm summer rain events and/or 40 kg N ha^?1 yr^?1. Increased summer precipitation and N additions did not have an overall significant effect on any of the measured root parameters. However, differences in winter precipitation resulting from interannual variability in rainfall appeared to affect root parameters with root production and turnover increasing following a wet winter most likely due to stimulation of annual grasses. In addition, roots were distributed more deeply in the soil following the wet winter. Root length density was initially higher under canopies compared to canopy interspaces, but converged toward the end of the study. In addition, roots tended to be distributed more deeply into the soil in canopy interspace areas. Results from this study indicated that increased summer precipitation and N deposition in response to climate change and urbanization are not likely to affect fine root dynamics in these Mojave Desert ecosystems, despite studies showing aboveground plant physiological responses to these environmental perturbations. However, changes in the amount and possibly distribution of winter precipitation may affect fine root dynamics.     

The distribution of tree and grass roots in savannas in relation to soil nitrogen and water

Here we describe the fine root distribution of trees and grasses relative to soil nitrogen and water profiles. The primary objective is to improve our understanding of edaphic processes influencing the relative abundance of trees and grasses in savanna systems. We do this at both a mesic (737 mm MAP) site on sandy-loam soils and at an arid (547 mm MAP) site on clay rich soils in the Kruger National Park in South Africa. The proportion of tree and grass fine roots at each soil depth were estimated using the δ¹³C values of fine roots and the δ¹³C end members of the fine roots of the dominant trees and grasses at our study sites. Changes in soil nitrogen concentrations with depth were indexed using total soil nitrogen concentrations and soil δ¹⁵N values. Soil water content was measured at different depths using capacitance probes. We show that most tree and grass roots are located in the upper layers of the soil and that both tree and grass roots are present at the bottom of the profile. We demonstrate that root density is positively related to the distribution of soil nitrogen and negatively related to soil moisture. We attribute the negative correlation with soil moisture to evaporation from the soil surface and uptake by roots. Our data is a snapshot of a dynamic process, here the picture it provides is potentially misleading. To understand whether roots in this system are primarily foraging for water or for nitrogen future studies need to include a dynamic component.     

Effect of Nitrogen, Phosphorus and Potassium Additions on Plant Biomass and Soil Nutrient Content of a Swale Barrier Strand Community in Louisiana

Fertilization of a swale plant community with various levelsand combinations of nitrogen, phosphorus and potassium resultedin increased plant growth. Nitrogen addition produced the greatestincrease in biomass. At some high nitrogen levels, phosphoruslimited plant production. Potassium fertilization did not influenceplant growth. Phosphorus fertilization decreased the levelsof soil calcium, magnesium and manganese. A shore-perpendiculargradient in apparent saltwater inundation influenced soil pHand the concentrations of iron, phosphorus, and magnesium. Fertilizer effects, swale Scirpus americanus, Spartina patens, nitrogen limitation     

Soil Moisture Rather than Soil Nutrient Regulates the Belowground Bud Bank of Rhizomatous Species <i>Psammochloa villosa</i> in Arid Sand Dunes

In arid and semi-arid sand dune ecosystems, belowground bud bank plays an important role in population regeneration and vegetation restoration. However, the responses of belowground bud bank size and composition to sand burial and its induced changes in soil environmental factors have been rarely studied. In arid sand dunes of Northwestern China, we investigated belowground bud bank size and composition of the typical rhizomatous psammophyte Psammochloa villosa as well as three key soil environmental factors (soil moisture, total carbon and total nitrogen) under different depths of sand burial. Total buds and rhizome buds increased significantly with increasing burial depth, whereas tiller buds first increased and then decreased, with a peak value at the depth of 20–30 cm. Soil moisture increased significantly with sand burial depth, and was positively correlated with the number of all buds and rhizome buds. Soil total carbon concentration first increased and then decreased with sand burial depth, and total nitrogen concentration was significantly lower under deep sand burial than those at shallow depths, and only the number of tiller buds was positively correlated with soil total nitrogen concentration. These results indicate that soil moisture rather than soil nutrient might regulate the belowground bud bank of P. villosa, and that clonal psammophytes could regulate their belowground bud bank in response to sand burial and the most important environmental stress (i.e., soil moisture). These responses, as the key adaptive strategy, may ensure clonal plant population regeneration and vegetation restoration in arid sand dunes.