Root morphological plasticity and nutrient acquisition of perennial grass species from habitats of different nutrient availability

We studied the root foraging ability and its consequences for the nutrient acquisition of five grass species that differ in relative growth rate and that occur in habitats that differ widely in nutrient availability. Foraging responses were quantified, based on the performance of the plants in homogeneous and heterogeneous soil environments of the same overall nutrient availability. Although all species tended to produce a significantly higher root length density in a nutrient-rich patch, this response was significant only for the faster-growing species. The increased root length density resulted from small, though not significant, changes in root biomass and specific root length. The effectiveness of root proliferation was determined by quantifying the total amount of nutrients (N and P) accumulated by the plants over the course of the experiment. Plants acquired more N in a heterogeneous environment than in a homogeneous environment, although the total nutrient availability was the same. The ability to acquire nutrients (N or P) in the heterogeneous environment was not related to the ability of species to increase root length density in response to local nutrient enrichment. In contrast to other studies, our results suggest that the role of morphological plasticity of roots in acquiring patchily distributed resources is limited. Possible reasons for this discrepancy are discussed. Received: 11 September 1997 / Accepted: 28 February 1998     

Growth and morphological responses to water level and nutrient supply in three emergent macrophyte species

Sanjiang Plain is the largest freshwater marsh in China, where plant zonation along water-level gradients is a common phenomenon. The aim of this experiment was to identify the role of water level and nutrient availability on plant zonation in the plain. Growth and root morphology of three perennial emergent macrophyte species were investigated by growing in two water levels (0.1 and 10.0 cm, relative to soil surface) and in two levels of nutrient supply (0 and 0.5 g slow-release fertilizer per container). In the plain, Carex lasiocarpa typically occurs at low elevations, Glyceria spiculosa at medial elevations, and Deyeuxia angustifolia at high elevations. The relative growth rate was the highest in C. lasiocarpa and the lowest in D. angustifolia in the 10.0-cm water level. Among the three species, only total biomass of D. angustifolia was affected by water level, and decreased with increasing water level. High nutrient supply led to increased total biomass in C. lasiocarpa and G. spiculosa. High water level led to an increased root diameter in G. spiculosa and a decreased root length in C. lasiocarpa. In the 10.0-cm water level, low nutrient supply led to thinner roots in D. angustifolia, but resulted in an increased specific root length (SRL) in C. lasiocarpa and root diameter in G. spiculosa. Water-level effect on root porosity was only observed in G. spiculosa, and nutrient amendment did not influence root porosity in all the species. These data indicate that both nutrient and water level are important factors regulating plant distribution pattern in the Sanjiang Plain, because both C. lasiocarpa and G. spiculosa are relatively sensitive to nutrient supply whereas D. angustifolia is sensitive to water level. Handling editor: S. M. Thomaz     

Root growth and plant biomass in <Emphasis Type="Italic">Lolium perenne</Emphasis> exploring a nutrient-rich patch in soil

We investigated soil exploration by roots and plant growth in a heterogeneous environment to determine whether roots can selectively explore a nutrient-rich patch, and how nutrient heterogeneity affects biomass allocation and total biomass before a patch is reached. Lolium perenne L. plants were grown in a factorial experiment with combinations of fertilization (heterogeneous and homogeneous) and day of harvest (14, 28, 42, or 56 days after transplanting). The plant in the heterogeneous treatment was smaller in its mean total biomass, and allocated more biomass to roots. The distributions of root length and root biomass in the heterogeneous treatment did not favor the nutrient-rich patch, and did not correspond to the patchy distribution of inorganic nitrogen. Specific root length (length/biomass) was higher and root elongation was more extensive both laterally and vertically in the heterogeneous treatment. These characteristics may enable plants to acquire nutrients efficiently and increase the probability of encountering nutrient-rich patches in a heterogeneous soil. However, heterogeneity of soil nutrients would hold back plant growth before a patch was reached. Therefore, although no significant selective root placement in the nutrient-rich patch was observed, plant growth before reaching nutrient-rich patches differed between heterogeneous and homogeneous environments.     

Root foraging for patchy resources in eight herbaceous plant species

The root foraging strategy of a plant species can be characterized by measuring foraging scale, precision, and rate. Trade-offs among these traits have been predicted to contribute to coexistence of competitors. We tested for trade-offs among root foraging scale (total root mass and length of structural roots), precision (ln-ratio of root lengths in resource-rich and resource-poor patches), and rate (days required for roots to reach a resource-rich patch, or growth rate of roots within a resource-rich patch) in eight co-occurring species. We found that root foraging scale and precision were positively correlated, as were foraging scale and the rate of reaching patches. High relative growth rate of a species did not contribute to greater scale, precision, or rate of root foraging. Introduced species had greater foraging scale, precision, and rate than native species. The positive correlations between foraging scale and foraging precision and rate may give larger species a disproportionate advantage in competition for patchy soil resources, leading to size asymmetric competition below ground.     

Robustness of demographic estimates in studies of plant responses to fire

Abstract For demographic models of resprouting species in relation to fire regimes, spatially separated populations of different ages are usually treated as a chronosequence, and the average behaviour of these populations is used as the estimate of the generalized behaviour of a single population through time. Using the data of Bradstock (1990), we examine the way in which the between-population variability affects the demographic parameters for Banksia serrata (Proteaceae) and thus assess whether the estimates of the shortest inter-fire interval necessary to maintain a stable population size under low and high intensity fires are robust to variation in these parameters. The range of our estimates compares well to the average estimate of Bradstock (1990) for high intensity fires, but not for low intensity fires. However the relevant demographic parameters appear to vary more between populations than they do as a result of the length of the inter-fire intervals, and the specific demographic model discussed here may have only local application.     

Fine root responses to temporal nutrient heterogeneity and competition in seedlings of two tree species with different rooting strategies

There is little direct evidence for effects of soil heterogeneity and root plasticity on the competitive interactions among plants. In this study, we experimentally examined the impacts of temporal nutrient heterogeneity on root growth and interactions between two plant species with very different rooting strategies: Liquidambar styraciflua (sweet gum), which shows high root plasticity in response to soil nutrient heterogeneity, and Pinus taeda (loblolly pine), a species with less plastic roots. Seedlings of the two species were grown in sandboxes in inter‐ and intraspecific combinations. Nutrients were applied in a patch either in a stable (slow‐release) or in a variable (pulse) manner. Plant aboveground biomass, fine root mass, root allocation between nutrient patch and outside the patch, and root vertical distribution were measured. L. styraciflua grew more aboveground (40% and 27% in stable and variable nutrient treatment, respectively) and fine roots (41% and 8% in stable and variable nutrient treatment, respectively) when competing with P. taeda than when competing with a conspecific individual, but the growth of P. taeda was not changed by competition from L. styraciflua. Temporal variation in patch nutrient level had little effect on the species’ competitive interactions. The more flexible L. styraciflua changed its vertical distribution of fine roots in response to competition from P. taeda, growing more roots in deeper soil layers compared to its roots in conspecific competition, leading to niche differentiation between the species, while the fine root distribution of P. taeda remained unchanged across all treatments. Synthesis. L. styraciflua showed greater flexibility in root growth by changing its root vertical distribution and occupying space of not occupied by P. taeda. This flexibility gave L. styraciflua an advantage in interspecific competition.     

Cd对不同种类植物生长和养分积累的影响

溶液培养法研究了不同活度Cd2 对4种植物卷心菜、黑麦草、玉米和白三叶草生长和养分积累的影响．结果表明,随Cd2 活度增加,4种植物的生长速率和干物质产量均下降．白三叶草对Cd2 最敏感;黑麦草和玉米耐性较强;卷心菜在高ACd2 时敏感．Cd2 对Fe、Mn、Cu、Zn、Ca、Mg、P、S等养分积累的影响因植物种类而异．Cd2 降低Fe、Mn、Cu、Zn、Ca、Mg等的积累．P在4种植物中和S在除卷心菜外的3种植物中的积累比对照增加．卷心菜和白三叶草对Cd毒敏感性差异与Fe、Mn、Ca、Mg的积累受Cd2 影响程度不同密切相关．     

Growth rate, root development and nutrient uptake of 55 plant species from the Great Plains Grasslands, USA

Empirical and theoretical studies have highlighted that plantcompetition and species diversity are substantially affected by interactionsamong plant growth and nutrient uptake rates, root lateral spread, rootplasticity, and small scale soil nutrient heterogeneity. This study wasdesignedto (a) experimentally estimate parameters regarding rootscaling patterns, root biomass allocation, growth rates, nutrient productivity,and root nutrient influx rates of 55 plant species common to Great Plainsgrasslands; and (b) determine if grasses and forbs can beclassified into statistically distinct groups based on these characteristics.Wefound that: (1) In all species root lateral spread, rootlength, and root surface area had significant allometric scaling relationshipswith root biomass, but that the relationships were unaffected by Navailability.(2) Reductions in the supply of N increased the root:shootratio in 62% of the species. (3) The frequency distributionandmean values of maximum relative growth rates were very similar for grasses andforbs/shrubs, but mid successional grasses had a higher relative growth ratethan late successional ones. (4) In 78% of the species tested,N productivity was increased by reductions in the N supply.(5)When subjected to a high N supply, the N and P productivity of grasses was, onaverage, higher than that of forbs/shrubs, and the N and P productivity ofC₄ grasses was, on average, higher than that of C₃grasses. No differences were found under a low N supply. (6)Nodifferences on the average maximum N and P influx rates per unit of rootsurfacearea were found between grasses and forbs or between C₃ andC₄ grasses, but both were correlated with maximum relative growthrate. (7) The set of parameters we measured were able toseparate grasses and forbs/shrubs into statistically distinct groups that tendto follow in broad terms the coarse vs. fine scaleforaging strategies hypothesis.     

镧对水稻根质膜透性和根分泌物中几种营养离子含量的影响

通过水培实验,研究了系列浓度La在不同处理时间内对水稻根质膜透性和根分泌物中几种营养离子含量的影响。结果表明,La在低浓度时(≤50μg·ml^-1),可以稳定细胞膜,减少电解质外渗率,在此浓度下,根系分泌物中K^+、Ca^2+、P和H^+的含量比对照低,La的浓度升高(100～400μg·ml^-1),随着处理时间的延长,电解质外渗率呈现短时间抑制,长时间促进的规律．La在高浓度时(c≥500μg·ml^-1),即使短时间的处理,也会使根细胞原生质膜破坏,电解质外渗率提高．此时根分泌物中K^+、Ca^2+、P和H^+含量均比对照高．由此可以得出在水培条件下,La^3+≤50μg·ml^-1对水稻生长是安全的。     

Spatial heterogeneity in species composition constrains plant community responses to herbivory and fertilisation

Environmental change can result in substantial shifts in community composition. The associated immigration and extinction events are likely constrained by the spatial distribution of species. Still, studies on environmental change typically quantify biotic responses at single spatial (time series within a single plot) or temporal (spatial beta diversity at single time points) scales, ignoring their potential interdependence. Here, we use data from a global network of grassland experiments to determine how turnover responses to two major forms of environmental change – fertilisation and herbivore loss – are affected by species pool size and spatial compositional heterogeneity. Fertilisation led to higher rates of local extinction, whereas turnover in herbivore exclusion plots was driven by species replacement. Overall, sites with more spatially heterogeneous composition showed significantly higher rates of annual turnover, independent of species pool size and treatment. Taking into account spatial biodiversity aspects will therefore improve our understanding of consequences of global and anthropogenic change on community dynamics.     

Increased nutrient supply facilitates acclimation to high-water level in the marsh plant Deyeuxia angustifolia: The response of root morphology

Both water level and nutrient availability are important factors influencing the growth of wetland plants. Increased nutrient supply might counteract the negative effects of flooding on the growth of the fast-growing species. Experimental evidence is scarce and the mechanism is far from clear. The aim of this study is to identify the role of nutrient availability in acclimation to high-water level by investigating the growth and root morphology of the marsh plant Deyeuxia angustifolia, one of the dominant species in the Sanjiang Plain, China. Experimental treatments included two water levels (0 and 10 cm, relative to soil surface) and three levels of nutrient supply (0, 0.5 and 1 g fertilizer per container). High-water level usually led to decreased biomass accumulation, shoot mass and root mass, whereas biomass accumulation was unaffected by water level at the highest nutrient level, indicating that high-nutrient availability played a role in compensating for the growth loss induced by the high-water level. Increased nutrient supply led to decreased root length in 0 cm water-level treatments, but root length increased with nutrient supply in the 10 cm water-level treatments. High-water level usually led to a lower lateral root density, lateral root:main root length ratio and the diameter of main roots and laterals, whereas increased nutrient supply resulted in thicker main roots or laterals, and a higher total root length, lateral root density and lateral root:main root length ratio. These data indicate that the growth of D. angustifolia is restrained by high-water level, and that increased nutrient supply not only ameliorates root characteristics to acclimate to high-water level but also results in a high-total root length to facilitate nutrient acquisition.     

Plant demographic responses to mycorrhizal symbiosis in tallgrass prairie

The effects of mycorrhizal symbiosis on seedling emergence, flowering and densities of several grasses and forbs were assessed in native tallgrass prairie and in sown garden populations at the Konza Prairie in northeastern Kansas. Mycorrhizal activity was experimentally suppressed with the fungicide benomyl. Flowering and stem densities of the cool-season grass, Dichanthelium oligosanthes, sedges (Carex spp.), and the forb Aster ericoides were higher in non-mycorrhizal (benomyl-treated) than in mycorrhizal plots and the magnitude of these differences was significantly affected by burning. Mycorrhizae significantly enhanced flowering of the warmseason grasses Andropogon gerardii and Sorghastrum nutans in burned prairie, but not in unburned sites. These patterns suggest that mycorrhizal effects on the dynamics of cool-season graminoid and forb populations are likely to be mediated indirectly through effects of the symbiosis on the competitive dominance of their neighbors. Seedling emergence rates of the cool-season C₃ grasses Elymus canadensis and Koeleria cristata were significantly reduced in the benomyl-treated plots, whereas benomyl treatment had no significant effect on seedling emergence of the warm-season C₄ grasses A. gerardii and Panicum virgatum. The forbs showed variable responses. Seedling emergence of Liatris aspera was greater under mycorrhizal conditions, but that of Dalea purpurea was unaffected by mycorrhizal treatment. These results show that effects of mycorrhizal symbiosis on the population dynamics of co-occurring prairie plants vary significantly both among species and among different life history stages within species. The results also indicate that mycorrhizas and fire interact to influence competitive interactions and demographic patterns of tallgrass prairie plant populations.     

Shifts in plant nutrient use strategies under secondary forest succession

In evergreen broad-leaved forests (EBLFs) in Tiantong National Forest Park, Eastern China, we studied the soil chemistry and plant leaf nutrient concentration along a chronosequence of secondary forest succession. Soil total N, P and leaf N, P concentration of the most abundant plant species increased with forest succession. We further examined leaf lifespan, leaf nutrient characteristics and root–shoot attributes of Pinus massoniana Lamb, the early-successional species, Schima superba Gardn. et Champ, the mid-successional species, and Castanopsis fargesii Franch, the late-successional species. These species showed both intraspecific and interspecific variability along succession. Leaf N concentration of the three dominant species increased while N resorption tended to decrease with succession; leaf P and P resorption didn’t show a consistent trend along forest succession. Compared with the other two species, C. fargesii had the shortest leaf lifespan, largest decay rate and the highest taproot diameter to shoot base diameter ratio while P. massoniana had the highest root–shoot biomass ratio and taproot length to shoot height ratio. Overall, P. massoniana used ‘conservative consumption’ nutrient use strategy in the infertile soil conditions while C. fargesii took up nutrients in the way of ‘resource spending’ when nutrient supply increased. The attributes of S. superba were intermediate between the other two species, which may contribute to its coexistence with other species in a wide range of soil conditions.     

Shading reduces exploitation of soil nitrate and phosphate by Agropyron desertorum and Artemisia tridentata from soils with patchy and uniform nutrient distributions

 Shading may both lessen the demand for soil nutrients and also the energy supply for nutrient acquisition. Since root foraging for nutrients in patchy environments can be energy-costly, especially for an immobile nutrient such as phosphate (P), the effects of shading may be most expected in heterogeneous soils. Plant acquisition of nitrate (N) and phosphate from soils with patchy and uniform nutrient distributions was determined in a field study under open sunlight and with shading for two common perennial Great Basin shrub steppe species, Agropyron desertorum and Artemisia tridentata. Partial shading in a pattern which can occur in shrub steppe vegetation significantly decreased plant N and P acquisition from soils both in the patchy and the uniform nutrient treatments. Artemisia was more affected by the shading than was Agropyron. Exploitation of the rather immobile P ion by both species was reduced to a much greater degree by the shading in the patchy distribution treatment than in the uniform nutrient treatment. As expected, plant acquisition of the more mobile N varied little with nutrient distribution treatment for both species and the depression of N acquisition by shading was the same in both nutrient distributions. The effects of shading appeared to have had its primary influence on different components of root foraging in the two species, especially in the nutrient-rich patches. For Agropyron shading primarily affected root proliferation, as indicated by reduced root density in patches. For Artemisia, shading most influenced root physiological uptake capacity and this was most pronounced in the nutrient-rich patches. While aboveground competition for light may generally reduce nutrient acquisition, the effects appear to be most pronounced if root systems of these steppe species are foraging for nutrients such as P in spatially heterogeneous soils. Received: 29 February 1996 / Accepted: 16 July 1996     

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.     

Root CEC determinations to establish root biomasses of two plant species grown in mixtures

Summary The use of measurements on the cation exchange capacity (CEC) of roots in competition experiments with two plant species is suggested. If the root-CEC differences between the two species are not too small, measurement of this parameter offers a very simple way to determine the separate contributions of the two plant species in the total root biomass of the mixture. This root-CEC method can be applied to total root yields as well as to detailed root-zone studies.Grassland Species Research Group, Publication no. 48     

Plant defensins and virally encoded fungal toxin KP4 inhibit plant root growth

Plant defensins are small, highly stable, cysteine-rich antimicrobial proteins that are thought to constitute an important component of plant defense against fungal pathogens. There are a number of such defensins expressed in various plant tissues with differing antifungal activity and spectrum. Relatively little is known about the modes of action and biological roles of these proteins. Our previous work on a virally encoded fungal toxin, KP4, from Ustilago maydis and subsequently with the plant defensin, MsDef1, from Medicago sativa demonstrated that some of these proteins specifically blocked calcium channels in both fungi and animals. The results presented here demonstrate that KP4 and three plant defensins, MsDef1, MtDef2, and RsAFP2, all inhibit root growth in germinating Arabidopsis seeds at low micromolar concentrations. We have previously demonstrated that a fusion protein composed of Rab GTPase (RabA4b) and enhanced yellow fluorescent protein (EYFP) is dependent upon calcium gradients for localization to the tips of the growing root hairs in Arabidopsis thaliana. Using this tip-localized fusion protein, we demonstrate that all four proteins rapidly depolarize the growing root hair and block growth in a reversible manner. This inhibitory activity on root and root hair is not directly correlated with the antifungal activity of these proteins and suggests that plants apparently express targets for these antifungal proteins. The data presented here suggest that plant defensins may have roles in regulating plant growth and development. A. Allen and A.K. Snyder contributed equally.