Co-occurring graminoid and forb species do not differ in their root morphological response to soil heterogeneity

We measured the proliferation of roots into experimental nutrient patches in a grassland community, distinguishing roots of graminoids and forbs. Biomass, length, and specific length were estimated for roots of each of the two functional groups, collected from patches differing in nutrient concentration, and established at four different times during a season. The ratio of graminoid and forb roots was compared with the graminoid-forb ratio in the above-ground biomass. Plant roots proliferated more intensively into patches with higher nutrient concentration, but the roots of the two functional groups had a similar ability to target richer patches. Relative proportion of graminoids was higher below-ground than above-ground and changed during the season, being lowest after mowing. Specific root length was higher for graminoid species, but did not respond to nutrient concentration in patches for either functional group. This is the first study to provide comparative information about root morphological response for graminoids and forbs, measured in a real, semi-natural plant community. The study shows no significant overall difference in the ability of these two functional types to place roots into nutrient-rich patches, but indicates other important differences among the two functional groups.     

Invasive forbs differ functionally from native graminoids, but are similar to native forbs

? Exotic plant invasions can alter ecosystem processes, particularly if the invasive species are functionally different from native species. We investigated whether such alterations can be explained by differences in functional traits between native and invasive plants of the same functional group or by differences in functional group affiliation. ? We compared six invasive forbs in Europe with six native forbs and six native graminoids in leaf and whole-plant traits, plasticity in response to nutrient supply and interspecific competition, litter decomposition rate, effects on soil nutrient availability, and allelopathy. All traits were measured in a series of pot experiments, and leaf traits additionally in the field. ? Invasive forbs differed from native forbs for only a few traits; they had less leaf chlorophyll and lower phosphorus (P) uptake from soil, but they tended to have a stronger allelopathic effect. The invasive forbs differed in many traits from the native graminoids, their leaves had lower tissue densities and a shorter life span, their litter decomposed faster and they had a lower nitrogen-use efficiency. ? Our results suggest that invasive forbs have the potential to alter ecosystem properties when invading graminoid-dominated and displacing native graminoids but not when displacing native forbs.     

Effects of Soil Nutrient Heterogeneity and Earthworms on Aboveground Biomass of Experimental Plant Communities

Soil nutrients are commonly heterogeneously distributed and earthworms are one of the most common soil organisms. While effects of both soil nutrient heterogeneity and earthworms have been well studied, their interactive effect on plant community productivity has rarely been tested. In a greenhouse experiment, we constructed experimental plant communities by sowing seed mixtures of four grasses, two legumes and two forbs in either a heterogeneous soil consisting of low and high nutrient soil patches or a homogeneous soil where the low and high nutrient soil patches were evenly mixed. The earthworm Eisenia fetida was either added to these soils or not. Aboveground biomass of the whole communities, grasses and legumes did not differ between the homogeneous and heterogeneous soils or between the soils with and without earthworms. However, soil nutrient heterogeneity reduced aboveground biomass of forbs, and such an effect did not interact with earthworms. In response to soil heterogeneity and earthworms, biomass ratio of the three functional groups showed similar patterns as that of their biomass. At the patch level, aboveground biomass of the whole community, grasses and legumes were greater in the high than in the low nutrient soil patches within the heterogeneous soil. A similar pattern was found for the forbs, but this was only true in the absence of earthworms. Our results suggest that soil nutrient heterogeneity and earthworms may not influence aboveground biomass of plant communities, despite the fact that they may modify the growth of certain plant functional groups within the community.     

Foliar macronutrient concentrations of forest understorey species in relation to Ellenberg's indices and potential relative growth rate

In order to investigate broad patterns of variation of the foliar mineral nutrient concentrations of herbaceous plant communities in the ground layer of W Europe forests, correlations were examined between Ellenberg's indices (N-index: mineral nitrogen availability, R-index: pH, F-index: soil moisture and L-index: light intensity) and literature values of macronutrient concentrations for 84 forbs and 39 graminoid species. Significant, positive correlations were found between the R-index and the plants' concentration of Ca and K (forbs only) and between the N-index and the plants' concentration of K, P (forbs only) and N (forbs and graminoids). Multiple regressions showed that the N-index was the best predictor of the plants' concentration of N (forbs and graminoids), P (forbs) and Ca (graminoids) and the R-index of the plants' concentration of Ca and K (forbs). The mineral nutrient concentrations of graminoids were lower and less strongly correlated with Ellenberg's indices than those of forbs. It is argued that the mineral nutrient concentrations in the plants match the availability of mineral nutrients in the soil for N, P (N-index) and Ca (R-index), but not for K and Mg. Significant, positive correlations were found between potential relative growth rate and the concentration of some elements (N, P, K and Ca in forbs, Ca and Mg in graminoids). This suggests that the increase in the concentration of these elements in plants along fertility gradients is due, at least partly, to genetically controlled alterations of leaf anatomy associated with increasing potential relative growth rate.     

Root plasticity of native and invasive Great Basin species in response to soil nitrogen heterogeneity

Soil nutrients are heterogeneously distributed in natural systems. While many species respond to this heterogeneity through root system plasticity, little is known about how the magnitude of these responses may vary between native and invasive species. We quantified root morphological and physiological plasticity of co-occurring native and invasive Great Basin species in response to soil nitrogen heterogeneity and determined if trade-offs exist between these foraging responses and species relative growth rate or root system biomass. The nine study species included three perennial bunchgrasses, three perennial forbs, and three invasive perennial forbs. The plants were grown in large pots outdoors. Once a week for 4 weeks equal amounts of ¹⁵NH₄ ¹⁵NO₃ were distributed in the soil either evenly through the soil profile, in four patches, or in two patches. All species acquired more N in patches compared to when N was applied evenly through the soil profile. None of the species increased root length density in enriched patches compared to control patches but all species increased root N uptake rate in enriched patches. There was a positive relationship between N uptake rate, relative growth rate, and root system biomass. Path analysis indicated that these positive interrelationships among traits could provide one explanation of how invasive forbs were able to capture 2 and 15-fold more N from enriched patches compared to the native grasses and forbs, respectively. Results from this pot study suggest that plant traits related to nutrient capture in heterogeneous soil environments may be positively correlated which could potentially promote size-asymmetric competition belowground and facilitate the spread of invasive species. However, field experiments with plants in different neighbor environments ultimately are needed to determine if these positive relationships among traits influence competitive ability and invader success.     

Diversity loss, recruitment limitation, and ecosystem functioning: lessons learned from a removal experiment

A five-year removal experiment in which plant functional group diversity was manipulated found strong limitation of ecosystem functioning caused by the differing abilities of remaining functional groups to recruit into space left unoccupied by the plants removed. We manipulated functional group diversity and composition by removing all possible combinations of zero, one, or two plant functional groups (forbs, C₃ graminoids, and C₄ graminoids), as well as randomly chosen biomass at levels corresponding to the functional group removals, from a prairie grassland community. Although random biomass removal treatments showed no significant effect of removing biomass in general on ecosystem functions measured ( P >0.05), the loss of particular functional groups led to significant differences in above- ( P <0.001) and belowground ( P <0.001) biomass, rooting-zone ( P =0.001) and leached ( P =0.01) nitrogen, nitrogen mineralization ( P <0.001), and community drought resistance ( P =0.002). Many of these differences stemmed from the marked difference in the ways remaining functional groups responded to the experimental removals. Strong recruitment limitation of C₄ graminoids resulted in large areas of open ground, high nutrient leaching, and high community drought resistance in plots containing just this functional group. In contrast, rhizomatous C₃ graminoids quickly colonized space and used soil resources made available by the removal of other groups, leading to lower soil nitrate in plots containing C₃ graminoids. These effects of recruitment limitation on ecosystem functioning illustrate possible effects of diversity loss not captured by synthetic experiments in which diversity gradients are created by adding high densities of seeds to bare soil.     

Ectomycorrhizal symbiosis can enhance plant nutrition through improved access to discrete organic nutrient patches of high resource quality

It is known that roots can respond to patches of fertility; however, root proliferation is often too slow to exploit resources fully, and organic nutrient patches may be broken down and leached, immobilized or chemically fixed before they are invaded by the root system. The ability of fungal hyphae to exploit resource patches is far greater than that of roots due to their innate physiological and morphological plasticity, which allows comprehensive exploration and rapid colonization of resource patches in soils. The fungal symbionts of ectomycorrhizal plants excrete significant quantities of enzymes such as chitinases, phosphatases and proteases. These might allow the organic residue to be tapped directly for nutrients such as N and P. Pot experiments conducted with nutrient-stressed ectomycorrhizal and control willow plants showed that when high quality organic nutrient patches were added, they were colonized rapidly by the ectomycorrhizal mycelium. These established willows (0.5 m tall) were colonized by Hebeloma syrjense P. Karst. for 1 year prior to nutrient patch addition. Within days after patch addition, colour changes in the leaves of the mycorrhizal plants (reflecting improved nutrition) were apparent, and after I month the concentration of N and P in the foliage of mycorrhizal plants was significantly greater than that in non-mycorrhizal plants subject to the same nutrient addition. It seems likely that the mycorrhizal plants were able to compete effectively with the wider soil microbiota and tap directly into the high quality organic resource patch via their extra-radical mycelium. We hypothesize that ectomycorrhizal plants may reclaim some of the N and P invested in seed production by direct recycling from failed seeds in the soil. The rapid exploitation of similar discrete, transient, high-quality nutrient patches may have led to underestimations when determining the nutritional benefits of ectomycorrhizal colonization.     

土壤养分空间异质性与植物根系的觅食反应

植物在长期进化过程中,为了最大限度地获取土壤资源,对养分的空间异质性产生各种可塑性反应．包括形态可塑性、生理可塑性、菌根可塑性等．许多植物种的根系在养分丰富的斑块中大量增生,增生程度种间差异较大,并受斑块属性(斑块大小、养分浓度)、营养元素种类和养分总体供应状况的影响．植物还通过调整富养斑块中细根的直径、分枝角、节问距以及空间构型来实现斑块养分的高效利用．根系的生理可塑性及菌根可塑性可能在一定程度上影响其形态可塑性．生理可塑性表现为处于不同养分斑块上的根系迅速调整其养分吸收速率,从而增加单位根系的养分吸收,对在时间上和空间上变化频繁的空间异质性土壤养分的利用具有重要意义,可在一定程度上弥补根系增生反应的不足．菌根可塑性目前研究较少,一些植物种的菌根代替细根实现在富养斑块中的增生．菌根增生的碳投入养分吸收效率较高、根系增生对增加养分吸收的作用较复杂,取决于养分离子在土壤中的移动性能以及是否存在竞争植物;对植物生长(竞争能力)的作用因种而异,一些敏感种由此获得生长效益,而其它一些植物种受影响较小．植物个体对土壤养分空间异质性反应能力和生长差异,影响其在群落中的地位和命运,最终影响群落组成及其结构．     

Controls of primary productivity and nutrient cycling in a temperate grassland with year‐round production

Abstract Net primary production (NPP) and nutrient dynamics of grasslands are regulated by different biotic and abiotic factors, which may differentially affect functional plant groups. Most studies have dealt with grasslands that have extremely low or zero production over a significant period of the year. Here we explore the relative importance of a few environmental factors as controls of aerial and below‐ground plant biomass production and nutrient dynamics in a grassland that is active throughout the year. We investigate their effect on the response of three main plant functional groups (warm‐ and cool‐season graminoids and forbs). We conducted a factorial experiment in a continuously grazed site in the Flooding Pampa grassland (Argentina). Factors were seasons (summer, autumn, winter and spring), and environmental agents (mowing, shade, addition of phosphorus [P] and nitrogen [N]). N addition had the largest and most extended impact: it tripled aerial NPP in spring and summer but had no effect on below‐ground biomass. This positive effect was accompanied by higher N acquisition and higher soil N availability. Mowing increased aerial NPP in winter, increased root biomass in the first 10 cm during autumn and winter and promoted N and P uptake by plants. Shading did not affect aerial NPP, but stimulated N and P uptake by plants. P addition had no effect on aerial NPP, but increased shallow root biomass and its N content in spring, and tripled P accumulation in plant biomass. The three plant functional groups differentially accounted for these ecosystem‐level responses. Graminoids explained the greater biomass production of N‐fertilized plots and mowing tended to promote forbs. These results suggest that the environmental controls of aerial NPP in this grassland vary among seasons, differentially impact the major floristic groups, and affect the energy and nutrient transfer to herbivores.     

Small-scale spatial heterogeneity in the vertical distribution of soil nutrients has limited effects on the growth and development of Prosopis glandulosa seedlings

We examined the above- and below-ground responses of seedlings of the woody shrub Prosopis glandulosa to the spatial heterogeneity of soil nutrients within the root zone. We performed a microcosm experiment where seedlings were grown with different combinations of nutrients [nitrogen (N), phosphorus (P), and both combined (NP)] and under different levels of nutrient heterogeneity (nutrients supplied as patches located in the bottom and/or upper portion of rooting zone versus homogeneous distribution). Seedling morphology and biomass did not show a strong response to changes in nutrient ion or spatial heterogeneity. Height, number of leaves, and specific leaf area did not vary significantly between treatments. The number of leaves, foliar biomass, stem biomass and biomass allocation to stems of seedlings showed more responsiveness to the addition of N and NP than to the addition of P. The spatial heterogeneity of nutrients affected the diameter, root biomass and leaf N content. Seedlings had higher diameter and root biomass when the nutrients were homogeneously distributed as compared to their placement as patches in the bottom of the microcosms. Their leaf N concentration increased in those treatments where the nutrient patch was located in the lower half as compared to the upper half of the microcosms. Root foraging responses to nutrient patches varied with their location. Significant root proliferation was observed when patches of N, P and NP were located in the upper portion of the rooting zone; when they were located in the lower portion such a response was observed only for P. Despite our findings that Prosopis seedlings have a low overall responsiveness to small-scale vertical differences in soil nutrient heterogeneity, our results suggest that these differences may modify the growth dynamics of the secondary roots of this ecologically important dryland species during the early stages of its development.     

Patch dynamics in grazed subtropical native pastures in south‐east Queensland

Abstract Patch formation is common in grazed grasslands but the mechanisms involved in the formation and maintenance of patches are not clear. To increase our knowledge on this subject we examined possible reasons for patch formation and the influence of management on changes between patch states in three experiments in native pasture communities in the Crows Nest district, south‐east Queensland. In these communities, small‐scale patches (tall grassland (dominated by large and medium tussock grasses), short swards (dominated by short tussock grasses and sedges), and lawns (dominated by stoloniferous and/or rhizomatous grasses)) are readily apparent. We hypothesized that the formation of short sward and lawn patches in areas of tall grassland was due to combinations of grazing and soil fertility effects. This was tested in Experiment 1 by applying a factorial combination of defoliation, nutrient application and transplants of short tussock and stoloniferous species to a uniform area of tall grassland. Total species density declined during the experiment, was lower with high nutrient applications, but was not affected by defoliation. There were significant changes in abundance of species that provided support for our hypotheses. With light defoliation and low nutrients, the tall grassland remained dominated by large tussock grasses and contained considerable amounts of forbs. With heavy defoliation, the pastures were dominated by medium tussock grasses and there were significant decreases in forbs and increases in sedges (mainly with low nutrients) and stoloniferous grasses (mainly with high nutrients). Total germinable seed densities and those of most species groups were significantly lower in the heavy defoliation than the light defoliation plots. Total soil seed numbers were not affected by nutrient application but there were fewer seeds of the erect forbs and more sedge seeds in plots with high nutrients. The use of resting from grazing and fire to manage transitions between patches was tested. In Experiment 2 , changes in species density and abundance were measured for 5 years in the three patch types with and without grazing. Experiment 3 examined the effects of fire, grazing and resting on short sward patches over 4 years. In Experiment 2 , total species density was lower in lawn than short sward or tall grassland patches, and there were more species of erect forbs than other plant groups in all patch types. The lawn patches were originally dominated by Cynodon spp. This dominance continued with grazing but in ungrazed patches the abundance of Cynodon spp. declined and that of forbs increased. In the short sward patches, dominance of short tussock grasses continued with grazing but in ungrazed plots their abundance declined while that of large tussock grasses increased. The tall grassland patches remained dominated by large and medium tussock species. In Experiment 3 , fire had no effect on species abundance. On the grazed plots the short tussock grasses remained dominant but where the plots were rested from grazing the small tussock grasses declined and the large tussock grasses increased in abundance. The slow and relatively small changes in these experiments over 4 or 5 years showed how stable the composition of these pastures is, and that rapid changes between patch types are unlikely.     

Plant root response to heterogeneity of soil resources: Effects of nutrient patches, AM symbiosis, and species composition

Response of the underground parts of a whole grassland community to heterogeneity in soil resources was studied under field conditions. The increased availability of nutrients (N, P) in patches was combined with suppression of arbuscular mycorrhizal symbiosis (by fungicide benomyl) and with modification of substrate structure (sieved soil, sieved soil mixed with coarse or fine sand). Three experimental runs were established in 1999, each with seven replicate blocks. Root and rhizome biomass was determined before the experimental treatment and after 53 days of the experiment. Plant community composition in neighbourhood of patches was recorded. Nutrient application had the most pronounced effect on the root biomass grown into the patches (the root biomass increased 1.64 times). The root biomass in patches with fungicide application was 1.17 times higher than in the untreated patches. The modification of substrate structure had no effect on the root biomass in patches but it did modify the effect of benomyl. The amount of root biomass proliferating into the patches was not related to species composition of vegetation near the patches. However, higher frequency of grass tillers in the patch neighbourhood and the mycorrhizal status of neighbouring species correlated significantly with the amount of root biomass in the experimental patches. In the pre-treatment community, amount of root biomass was significantly related to the abundance of grasses and mycorrhizal forbs surrounding the sampling points.     

A dual porosity model of nutrient uptake by root hairs

? The importance of root hairs in the uptake of sparingly soluble nutrients is understood qualitatively, but not quantitatively, and this limits efforts to breed plants tolerant of nutrient-deficient soils. ? Here, we develop a mathematical model of nutrient uptake by root hairs allowing for hair geometry and the details of nutrient transport through soil, including diffusion within and between soil particles. We give illustrative results for phosphate uptake. ? Compared with conventional 'single porosity' models, this 'dual porosity' model predicts greater root uptake because more nutrient is available by slow release from within soil particles. Also the effect of soil moisture is less important with the dual porosity model because the effective volume available for diffusion in the soil is larger, and the predicted effects of hair length and density are different. ? Consistent with experimental observations, with the dual porosity model, increases in hair length give greater increases in uptake than increases in hair density per unit main root length. The effect of hair density is less in dry soil because the minimum concentration in solution for net influx is reached more rapidly. The effect of hair length is much less sensitive to soil moisture.     

Distribution of plant species at a biome transition zone in New Mexico

Question: Is there a difference in plant species and life form composition between two major patch types at a biome transition zone? Are subordinate species associated with different patch types at the shortgrass steppe — Chihuahuan desert grassland transition zone? Is this association related to differences in soil texture between patch types and the geographic range of associated species? Location: central New Mexico, USA. Methods: Patches dominated by either Bouteloua gracilis, the dominant species in the shortgrass steppe, or Bouteloua eriopoda, dominant species in the Chihuahuan desert grasslands, were sampled for the occurrence of subordinate species and soil texture within a 1500‐ha transitional mosaic of patches. Results: Of the 52 subordinate species analysed, 16 species were associated with B. gracilis‐dominated patches and 12 species with B. eriopoda‐dominated patches. Patches dominated by B. gracilis were richer in annual grasses and forbs, whereas patches dominated by B. eriopoda contained more perennials forbs and shrubs. Soils of B. gracilis‐dominated patches had higher clay and lower rock contents compared with soils of B. eriopoda‐dominated patches. Differences in species characteristics of the dominant species as well as differences in soil texture between patch types contribute to patch‐scale variation in composition. The association of species to patch types was not related to their geographic range and occurrence in the adjacent biomes. Conclusions: Patch types at this biome transition zone have characteristic life‐form and species composition, but species are associated to patch types due to local constraints, independently from their affinity to the adjacent biomes.     

Limited costs of wrong root placement in Rumex palustris in heterogeneous soils

Nutrient hot spots in the soil have a limited life span, but the costs and benefits for root foraging are still underexposed. We assessed short-term costs that may arise when a nutrient-rich patch induces root proliferation, but then rapidly disappears. Rumex palustris plants were grown with a homogeneous or a heterogeneous nutrient application. After root proliferation in a nutrient-rich patch, nutrient supply was switched from homogeneous to heterogeneous, and vice versa, or the patch location was changed. R. palustris proliferated its roots in the rich patch. After switching, the relative growth rates of the roots were adjusted to the novel pattern of nutrient availability. However, the changes in local root biomass lagged behind the rapid shift in nutrient supply, because the root mass realized in specific sectors could not be rapidly relocated. Despite this, R. palustris did not exhibit costs of switching in terms of biomass or nitrogen uptake. Our data suggest that rapid shifts in uptake rate and redistribution of nitrogen within the plant may have lowered the costs of incorrect root placement.     

Soil heterogeneity and community composition jointly influence grassland biomass

Question: Does the spatial pattern of nutrient supply modify community biomass responses to changes in both species composition and richness? Location: Duke University Phytotron (Durham, North Carolina, USA). Methods: We conducted a microcosm experiment to evaluate individual plant and whole community responses to species richness, species composition and soil nutrient heterogeneity. The experiment consisted of seven levels of species composition (all possible combinations of Lolium perenne, Poa pratensis and Plantago lanceolata) crossed with three levels of soil nutrient distribution (homogeneous, heterogeneous‐up, and heterogeneous‐down, where up and down indicates the location of a nutrient patch in either the upper or the lower half of the soil column, respectively). Results: Communities containing Plantago and Lolium responded to nutrient heterogeneity by increasing above‐ and below‐ground biomass. Nutrient heterogeneity also increased size inequalities among individuals of these species. Significant species composition X nutrient heterogeneity interactions on community biomass and individual size inequality were observed when nutrient patches were located in the upper 10 cm of the soil columns. However, root proliferation in nutrient patches was equivalent regardless of the vertical placement of the patch. Conclusions: Our results suggest that nutrient heterogeneity may interact with plant species composition to determine community biomass, and that small‐scale vertical differences in the location of nutrient patches affect individual and community responses to this heterogeneity.     

Biomass compensation and plant responses to 7 years of plant functional group removals

Question: What is the role of functional group identity in determining community composition and dynamics? Location: A natural grassland in Yukon Territory, Canada. Methods: We selectively removed single plant functional groups (graminoids, forbs, legumes) to examine their effects on biomass compensation, the distribution of biomass among common and rare colonizing species, and plant species richness and diversity. Removals were conducted across two environmental treatments (fertilization and fungicide) to test if biomass compensation was context‐dependent. Biomass was estimated non‐destructively using point‐intercept sampling. Results: When graminoids or legumes were continuously removed, there was full biomass compensation by the remaining functional groups after 5 years, but only partial compensation when forbs were removed. Biomass compensation depended on the colonizing functional group; forbs showed no increase in biomass until 5 years after the removal of any functional group, but graminoids colonized quickly after removals. After any removal, the dominant species within each remaining functional group showed no compensatory growth, whereas the first subdominant forb and graminoid both increased in biomass. Rare species had a delayed response to removals; rare species biomass only increased beginning 5 years after removals. Context dependence was observed only in the response of subdominant species to removals, and these responses did not translate into context‐dependent effects on total estimated biomass. Conclusion: We show that the effects of losing a plant functional group depends both on the identity of the group removed and on the species remaining. In this northern grassland, most compensatory growth was by the subdominant species, which may determine the direction of community development in the long term.     

Fine root growth and demographic responses to nutrient patches in four old-field plant species

Proliferation of roots in a nutrient patch can occur either as a result of an increase in root length (morphological response) or by a change in root birth or death rates (demographic responses). In this study we attempted to distinguish between these two mechanisms of response to nutrient patches and to compare the responses of four old-field plant species (two annuals, two perennials). For all four species combined, there were significant increases in root numbers and root length in fertilized patches. Root proliferation in fertilized patches was largely due to increased birth (=branching) rates of new roots. However, there was also a significant increase in root death rates in the fertilized patches which reduced the magnitude of the increase in net root numbers. Plots for individual species suggested they differed in the magnitude and timing of root proliferation in fertilized patches due to differences in root birth and death rates. However, because of the limited sample size in this study, there was only a marginally significant difference among species in root birth rates, and no difference in death rates. Further studies are currently underway to better quantify species differences in the demographic mechanism, as well as magnitude, of response to nutrient patches and if this would affect the ability to exploit small-scale heterogeneity in soil resources.     

Root responses to nutrient patches in grassland and forest

Differences between growth forms in root responses to experimentally created heterogeneity have been documented in many greenhouse and plot studies, but not in natural vegetation. Here we examined the response of roots to experimental nutrient patches in undisturbed grassland and forest at the northern edge of the North American Great Plains. Forest vegetation increases the spatial heterogeneity of soil resources, and we tested for differences between forest and grassland roots in response to patches. Ten minirhizotrons (clear tubes, 5 cm diameter, 180 cm long) were installed in both grassland and forest 3 years before the experiment. Minirhizotrons ran horizontally 10 cm beneath the soil surface. Patches of available nitrogen (N) were created over the tubes, using three concentrations (0, 3, 15 g N m⁻² yr⁻¹) and two patch sizes (1␣and 10 cm²). Root images were collected beneath patches over the course of a growing season. Root length was significantly greater in grassland than forest at the start and end of the growing season, but did not respond to N patches. Root production was also significantly greater in grassland than forest, and was significantly greater (about 20%) in high-N patches than in unfertilized patches. This increase, however, did not differ between vegetation types. Turnover did not vary with any treatment, and patch size had no effect on any response variable. Overall, differences caused by experimental patches were much smaller than differences between habitats, and did not vary between habitats. Realistic levels of experimentally imposed hetereogeneity in established vegetation may not be much greater than background levels, and field vegetation has extant root systems which respond to patches via uptake instead of growth. Both mechanisms should contribute to less root proliferation in field experiments than in greenhouse experiments.     

The timing and degree of root proliferation in fertile-soil microsites for three cold-desert perennials

Summary Root proliferation in nutrient-rich soil patches is an important mechanism facilitating nutrient capture by plants. Although the phenomenon of root proliferation is well documented, the specific timing of this proliferation has not been investigated. We studied the timing and degree of root proliferation for three perennial species common to the Great Basin region of North America: a shrub, Artemisia tridentata, a native tussock grass, Agropyron spicatum, and an introduced tussock grass, Agropyron desertorum. One day after we applied nutrient solution to small soil patches, the mean relative growth rate of Agropyron desertorum roots in these soil patches was two to four times greater than for roots of the same plants in soil patches reated with distilled water. Most of the increased root growth came from thin, laterally branching roots within the patches. This rapid and striking root proliferation by Agropyron desertorum occurred in response to N-P-K enrichment as well as to P or N enrichment alone. A less competitive bunchgrass, Agrophyron spicatum, showed no tendency to proliferate roots in enriched soil patches during these two-week experiments. The shrub Artemisia tridentata proliferated roots within one day of initial solution injection in the N-enrichment experiment, but root proliferation of this species was more gradual and less consistent in the N-P-K and P-enrichment experiments, respectively. The ability of Agropyron desertorum to proliferate roots rapidly may partly explain both its general competitive success and its superior ability to exploit soil nutrients compared to Agropyron spicatum in Great Basin rangelands of North America.