Substrate heterogeneities cause root growth trajectories to switch from vertical to oblique.

Hard pans, soil compaction, soil aggregation and stones create physical barriers that can affect the development of a root system. Roots are known to exploit paths of least resistance to avoid such obstacles, but the mechanism through which this is achieved is not well understood. Here, we combined 3D-printed substrates with a high-throughput live imaging platform to study the responses of plant roots to a range of physical barriers. Using image analysis algorithms, we determined the properties of growth trajectories and identified how the presence of rigid circular obstacles affects the ability of a primary root to maintain its vertical trajectory. Results showed the types of growth responses were limited, both vertical and oblique trajectories were found to be stable and influenced by the size of the obstacles. When obstacles were of intermediate sizes, trajectories were unstable and changed in nature through time. We formalised the conditions for root trajectory to change from vertical to oblique, linking the angle at which the root detaches from the obstacle to the root curvature due to gravitropism. Exploitation of paths of least resistance by a root may therefore be constrained by the ability of the root to curve and respond to gravitropic signals.

Heterogeneity promotes resilience in restored prairie: Implications for the environmental heterogeneity hypothesis.

Enhancing resilience in formerly degraded ecosystems is an important goal of restoration ecology. However, evidence for the recovery of resilience and its underlying mechanisms require long-term experiments and comparison with reference ecosystems. We used data from an experimental prairie restoration that featured long-term soil heterogeneity manipulations and data from two long-term experiments located in a comparable remnant (reference) prairie to (1) quantify the recovery of ecosystem functioning (i.e., productivity) relative to remnant prairie, (2) compare the resilience of restored and remnant prairies to a natural drought, and (3) test whether soil heterogeneity enhances resilience of restored prairie. We compared sensitivity and legacy effects between prairie types (remnant and restored) and among four prairie sites that included two remnant prairie sites and prairie restored under homogeneous and heterogeneous soil conditions. We measured sensitivity and resilience as the proportional change in aboveground net primary productivity (ANPP) during and following drought (sensitivity and legacy effects, respectively) relative to average ANPP based on 4 pre-drought years (2014-2017). In nondrought years, total ANPP was similar between remnant and restored prairie, but remnant prairie had higher grass productivity and lower forb productivity compared with restored prairie. These ANPP patterns generally persisted during drought. The sensitivity of total ANPP to drought was similar between restored and remnant prairie, but grasses in the restored prairie were more sensitive to drought. Post-drought legacy effects were more positive in the restored prairie, and we attributed this to the more positive and less variable legacy response of forb ANPP in the restored prairie, especially in the heterogeneous soil treatment. Our results suggest that productivity recovers in restored prairie and exhibits similar sensitivity to drought as in remnant prairie. Furthermore, creating heterogeneity promotes forb productivity and enhances restored prairie resilience to drought.

Site-Independent Mapping of Clay Content in Vineyard Soils via Mobile Proximal Gamma-Ray Spectrometry and Machine Learning Calibrations.

Vineyards hold considerable soil variability between regions and plots, and there is frequently large soil heterogeneity within plots. Clay content in vineyard soils is of interest with respect to soil management, environmental monitoring, and wine quality. However, spatially resolved clay mapping is laborious and expensive. Gamma-ray spectrometry (GS) is a suitable tool for predicting clay content in precision agriculture when locally calibrated, but it has scarcely been tested site-independently and in vineyards. This study evaluated GS to predict clay content with a site-independent calibration and four machine learning algorithms (Support Vector Machines, Random Forest, k-Nearest Neighbors, and Bayesian regulated neuronal networks) in eight vineyards from four German vine-growing regions. Clay content in the studied soils ranged from 62 to 647 g kg-1. The Random Forest calibration was most suitable. Test set evaluation revealed good model performance for the entire dataset with RPIQ = 4.64, RMSEP = 56.7 g kg-1, and R2 = 0.87; however, prediction quality varied between the sites. Overall, GS with the Random Forest model calibration was appropriate to predict the clay content and its spatial distribution, even for heterogeneous geopedological settings and in individual plots. Therefore, GS is considered a valuable tool for soil mapping in vineyards, where clay content and product quality are closely linked.

Multi-imaging platform for rhizosphere studies: Phosphorus and oxygen fluxes.

Rhizosphere is a soil volume of high spatio-temporal heterogeneity and intensive plant-soil-microbial interactions, for which visualization and process quantification is of highest scientific and applied relevance, but still very challenging. A novel methodology for quick assessment of two-dimensional distribution of available phosphorus (P) in rhizosphere was suggested, tested, and development up to the application platform. Available P was firstly trapped by an in-situ diffusive gradients in thin-films (DGT) sampler with precipitated zirconia as the binding gel, and subsequently, the loaded gel was analyzed with an optimized colorimetric imaging densitometry (CID). The imaging platform was established linking: i) DGT, ii) planar optode, and iii) soil zymography techniques to simultaneously determine available P, oxygen, and acid phosphatase in rhizosphere at sub-millimeter spatial scales. The DGT identified available P level in rice rhizosphere were spatially overlapping to the localized redox hotspots and phosphatase activity. The spatial relationship between available P and acid phosphatase activity was dependent on root development. The root radial oxygen loss (ROL) remained active during the experimental observations (2-3 days), while a flux of available P of 10 pg cm-2 s-1 was visualized within 2-3 mm of roots, confirming the correlative response of rice roots to oxygen secretion and P uptake. Summarizing, the established imaging platform is suitable to capture spatial heterogeneity and temporal dynamics of root activities, nutrient bioavailability, ROL and enzyme activities in rhizosphere.

Genotypic differences in systemic root responses to mechanical obstacles.

As roots grow through the soil to forage for water and nutrients, they encounter mechanical obstacles such as patches of dense soil and stones that locally impede root growth. Here, we investigated hitherto poorly understood systemic responses of roots to localised root impedance. Seedlings of two wheat genotypes were grown in hydroponics and exposed to impenetrable obstacles constraining the vertical growth of the primary or a single seminal root. We deployed high-resolution in vivo imaging to quantify temporal dynamics of root elongation rate, helical root movement, and root growth direction. The two genotypes exhibited distinctly different patterns of systemic responses to localised root impedance, suggesting different strategies to cope with obstacles, namely stress avoidance and stress tolerance. Shallower growth of unconstrained seminal roots and more pronounced helical movement of unconstrained primary and seminal roots upon localised root impedance characterised the avoidance strategy shown by one genotype. Stress tolerance to localised root impedance, as exhibited by the other genotype, was indicated by relatively fast elongation of primary roots and steeper seminal root growth. These different strategies highlight that the effects of mechanical obstacles on spatiotemporal root growth patterns can differ within species, which may have major implications for resource acquisition and whole-plant growth.

Multiple and contrasting pressures determine intraspecific phytochemical variation in a tropical shrub.

Intraspecific phytochemical variation across a landscape can cascade up trophic levels, potentially mediating the composition of entire insect communities. Surprisingly, we have little understanding of the processes that regulate and maintain phytochemical variation within species, likely because these processes are complex and operate simultaneously both temporally and spatially. To assess how phytochemistry varies within species, we tested the degree to which resource availability, contrasting soil type, and herbivory generate intraspecific chemical variation in growth and defense of the tropical shrub, Piper imperiale (Piperaceae). We quantified changes in both growth (e.g., nutritional protein, above- and below-ground biomass) and defense (e.g., imide chemicals) of individual plants using a well-replicated fully factorial shade-house experiment in Costa Rica. We found that plants grown in high light, nutrient- and richer old alluvial soil had increased biomass. High light was also important for increasing foliar protein. Thus, investment into growth was determined by resource availability and soil composition. Surprisingly, we found that chemical defenses decreased in response to herbivory. We also found that changes in plant protein were more plastic compared to plant defense, indicating that constitutive defenses may be relatively fixed, and thus an adaptation to chronic herbivory that is common in tropical forests. We demonstrate that intraspecific phytochemical variation of P. imperiale is shaped by resource availability from light and soil type. Because environmental heterogeneity occurs over small spatial scales (tens of meters), herbivores may be faced with a complex phytochemical landscape that may regulate how much damage any individual plant sustains.

Can plants integrate information on above-ground competition in their directional responses below ground?

BACKGROUND AND AIMS: Light competition can induce varying above-ground responses in plants. However, very little is known regarding the effect of above-ground light competition cues on plant responses below ground. Here we asked whether light competition cues that indicate the occurrence and direction of neighbours above ground might affect directional root placemat. METHODS: In a common-garden experiment, we examined the integrated responses of the annual procumbent plant Portulaca oleracea to light competition cues and soil nutrient distribution. Soil nutrients were distributed either uniformly or in patches, and light competition was simulated using a transparent green filter, which was spatially located either in the same or opposite direction of the soil nutrient patch. KEY RESULTS: As predicted, root proliferation of P. oleracea increased in the direction of the enriched soil patches but was homogenously distributed under the uniform nutrient distribution. Interestingly, root distribution was also affected by the light competition cue and increased in its direction regardless of the location of the soil patches. CONCLUSIONS: Our results provide initial support to the idea that below-ground plant responses to competition might also be regulated by above-ground neighbour cues, highlighting the need to further investigate the combined effects of both above- and below-ground competition cues on root behaviour.

Patterns of leaf trait variation underlie ecological differences among sympatric tree species of Damburneya in a tropical rainforest.

PREMISE: Although ecological differentiation driven by altitude and soil is hypothesized to promote coexistence of sympatric tree species of Damburneya (Lauraceae), the mechanistic role of leaf functional variation on ecological differentiation among co-occurring species remains unexplored. We aimed to determine whether the patterns of leaf trait variation reflect ecological differences among sympatric Damburneya species. We tested whether trait correlations underlying functional strategies and average species traits vary in response to local soil heterogeneity along an altitudinal gradient, potentially affecting species distributions. METHODS: At two contrasting altitudes (100, 1100 m a.s.l.) in a Mexican tropical rainforest, we characterized soil chemical and physical properties and sampled four Damburneya species to quantify five leaf functional traits. We used linear models to analyze paired and multivariate trait correlations, spatial and interspecific effects on trait variation, and trait response to local soil heterogeneity. Relative contributions of intra- and interspecific variation to local trait variability were quantified with an ANOVA. RESULTS: Soil nutrient availability was higher at low altitude, but all species had a high leaf N:P ratio across altitudes suggesting a limited P supply for plants. Species distribution differed altitudinally, with some species constrained to low or high altitude, potentially reflecting soil nutrient availability. Leaf traits responded to altitude and local soil properties, suggesting interspecific differences in functional strategies according to the leaf economics spectrum (conservative vs. acquisitive). CONCLUSIONS: The interspecific divergence in functional strategies in response to local environmental conditions suggests that trait variation could underlie ecological differentiation among Damburneya sympatric species.

PREMISA: Aunque se ha propuesto que la diferenciación ecológica impulsada por la variación del suelo y la altitud ha promovido la coexistencia de especies arbóreas simpátricas de Damburneya (Lauraceae), el papel de la variación de funcional foliar como mecanismo para explicar las diferencias ecológicas entre especies que coocurren sigue sin explorarse. Nuestro objetivo fue determinar si los patrones de variación de los rasgos funcionales foliares reflejan diferencias ecológicas entre especies simpátricas de Damburneya. Hipotetizamos que, tanto las correlaciones entre los rasgos que determinan las estrategias funcionales, como los valores promedio de los rasgos de las especies, varían en respuesta a la heterogeneidad local del suelo a lo largo de un gradiente altitudinal, afectando potencialmente la distribución de las especies. MÉTODO: En dos altitudes contrastantes (100, 1000 m s.n.m) en una selva húmeda tropical mexicana caracterizamos propiedades físicas y químicas del suelo y muestreamos cuatro especies de Damburneya para cuantificar cinco rasgos funcionales foliares. Usamos modelos lineales para analizar las correlaciones pareadas y multivariadas entre rasgos, los efectos espaciales e interespecíficos en la variación de rasgos, y la respuesta de los rasgos a la heterogeneidad local del suelo. Además, cuantificamos la contribución relativa de la variación intra- e interespecífica en la varianza local de los rasgos con un análisis de varianza. RESULTADOS: La disponibilidad de nutrientes en el suelo fue mayor en la zona baja, pero todas las especies tuvieron valores altos de N:P foliar en ambas altitudes, lo que sugiere un suministro limitado de P para las plantas. La distribución de las especies difirió altitudinalmente y algunas de ellas se restringieron a zonas altas o bajas, reflejando potencialmente la disponibilidad de nutrientes en el suelo. Los rasgos foliares respondieron a la altitud y a las propiedades edáficas locales, sugiriendo diferencias interespecíficas en las estrategias funcionales según el espectro de economía foliar (conservativa vs. adquisitiva). CONCLUSIONES: La divergencia interespecífica en las estrategias funcionales en respuesta a las condiciones ambientales locales sugiere que la variación de rasgos podría explicar algunas diferencias ecológicas entre las especies simpátricas de Damburneya.

Soil heterogeneity influence on the distribution of heavy metals in soil during acid rain infiltration: Experimental and numerical modeling.

Acid rain is a global environmental problem that mobilizes heavy metals in soils, while the distribution and geochemical fraction of heavy metals during acid rain infiltration in heterogeneous soils are still unclear. In this study, we performed column experiments to investigate the distribution and geochemical fraction of Cu, Pb, Ni and Cd in heterogeneously layered soils during continuous acid rain infiltration. Chloride ion used as a conservative tracer was found to be uniformly distributed during acid rain infiltration, showing insignificant preferential flow effects in the column. In contrast, however, the distribution of heavy metals was highly non-uniform, especially in the silty soil at the lower part of the column, indicating a heterogeneous distribution of adsorption capacity. In addition, in the control experiments with neutral rain infiltration, uniform distribution of heavy metals was observed, indicating that the heterogeneous distribution of adsorption coefficient during acid rain infiltration was mainly caused by different pH buffering capacities. A numerical model considering water flow and solute transport was developed, where the average water-solid distribution coefficient (Kd) in Layer 2 was only 1.5-12.5% of that in Layer 1 during acid rain infiltration. The model could predict the variation of heavy metal concentrations in soil with the majority of error less than 35%, confirming that different Kd induced the heterogeneous distribution of heavy metals. In addition, the geochemical fraction of heavy metals in the upper coarse sand layer remained stable, while the acid-extractable fractions in the lower loam and silt loam layer gradually increased. Our findings suggest that soil heterogeneity, especially chemical heterogeneity affected by rainfall acidity, has an important influence on the infiltration, migration and geochemical fraction of heavy metals in soils. This study could help guide the risk assessment of heavy metal-contaminated sites that were polluted by acid rain or landfill leachate.

Soybean Root Nodule and Rhizosphere Microbiome: Distribution of Rhizobial and Nonrhizobial Endophytes.

Soybean root nodules are known to contain a high diversity of both rhizobial endophytes and nonrhizobial endophytes (NREs). Nevertheless, the variation of these bacteria among different root nodules within single plants has not been reported. So far, it is unclear whether the selection of NREs among different root nodules within single plants is a random process or is strictly controlled by the host plant to favor a few specific NREs based on their beneficial influence on plant growth. As well, it is also unknown if the relative frequency of NREs within different root nodules is consistent or if it varies based on the location or size of a root nodule. We assessed the microbiomes of 193 individual soybean root nodules from nine plants using high-throughput DNA sequencing. Bradyrhizobium japonicum strains occurred in high abundance in all root nodules despite the presence of other soybean-compatible rhizobia, such as Ensifer, Mesorhizobium, and other species of Bradyrhizobium in soil. Nitrobacter and Tardiphaga were the two nonrhizobial genera that were uniformly detected within almost all root nodules, though they were in low abundance. DNA sequences related to other NREs that have frequently been reported, such as Bacillus, Pseudomonas, Flavobacterium, and Variovorax species, were detected in a few nodules. Unlike for Bradyrhizobium, the low abundance and inconsistent occurrence of previously reported NREs among different root nodules within single plants suggest that these microbes are not preferentially selected as endophytes by host plants and most likely play a limited part in plant growth as endophytes.IMPORTANCE Soybean (Glycine max L.) is a valuable food crop that also contributes significantly to soil nitrogen by developing a symbiotic association with nitrogen-fixing rhizobia. Bacterial endophytes (both rhizobial and nonrhizobial) are considered critical for the growth and resilience of the legume host. In the past, several studies have suggested that the selection of bacterial endophytes within root nodules can be influenced by factors such as soil pH, nutrient availability, host plant genotype, and bacterial diversity in soil. However, the influence of size or location of root nodules on the selection of bacterial endophytes within soybean roots is unknown. It is also unclear whether the selection of nonrhizobial endophytes within different root nodules of a single plant is a random process or is strictly regulated by the host. This information can be useful in identifying potential bacterial species for developing bioinoculants that can enhance plant growth and soil nitrogen.

An Evaluation of Different NIR-Spectral Pre-Treatments to Derive the Soil Parameters C and N of a Humus-Clay-Rich Soil.

Near-infrared reflectance spectroscopy (NIRS) was successfully used in this study to measure soil properties, mainly C and N, requiring spectral pre-treatments. Calculations in this evaluation were carried out using multivariate statistical procedures with preceding pre-treatment procedures of the spectral data. Such transformations could remove noise, highlight features, and extract essential wavelengths for quantitative predictions. This frequently significantly improved the predictions. Since selecting the appropriate transformation was not straightforward due to the large numbers of available methods, more comprehensive insight into choosing appropriate and optimized pre-treatments was required. Therefore, the objectives of this study were (i) to compare various pre-processing transformations of spectral data to determine their suitability for modeling soil C and N using NIR spectra (55 pre-treatment procedures were tested), and (ii) to determine which wavelengths were most important for the prediction of C and N. The investigations were carried out on an arable field in South Germany with a soil type of Calcaric Fluvic Relictigleyic Phaeozem (Epigeoabruptic and Pantoclayic), created in the flooding area of the Isar River. The best fit and highest model accuracy for the C (Ct, Corg, and Ccarb) and N models in the calibration and validation modes were achieved using derivations with Savitzky-Golay (SG). This enabled us to calculate the Ct, Corg, and N with an R2 higher than 0.98/0.86 and an ratio of performance to the interquartile range (RPIQ) higher than 10.9/4.1 (calibration/validation).

Comparison of plant-soil feedback experimental approaches for testing soil biotic interactions among ecosystems.

The study of interactions and feedbacks between plants and soils is a rapidly expanding research area, and a primary tool used in this field is to perform glasshouse experiments where soil biota are manipulated. Recently, there has been vigorous debate regarding the correctness of methods for carrying out these types of experiment, and specifically whether it is legitimate to mix soils from different sites or plots (mixed soil sampling, MSS) or not (independent soil sampling, ISS) to create either soil inoculum treatments or subjects. We performed the first empirical comparison of MSS vs ISS approaches by comparing growth of two boreal tree species (Picea abies and Pinus sylvestris) in soils originating from 10 sites near the boreal forest limit in northern Sweden, and 10 sites in the subarctic region where boreal forests may potentially expand as a result of climate change. We found no consistent differences in the conclusions that we reached whether we used MSS or ISS approaches. We propose that researchers should not choose a soil handling method based on arguments that one method is inherently more correct than the other, but rather that method choice should be based on correct alignment with specific research questions and goals.

Soil aggregates as biogeochemical reactors and implications for soil-atmosphere exchange of greenhouse gases-A concept.

Soil-atmosphere exchange significantly influences the global atmospheric abundances of carbon dioxide (CO2 ), methane (CH4 ), and nitrous oxide (N2 O). These greenhouse gases (GHGs) have been extensively studied at the soil profile level and extrapolated to coarser scales (regional and global). However, finer scale studies of soil aggregation have not received much attention, even though elucidating the GHG activities at the full spectrum of scales rather than just coarse levels is essential for reducing the large uncertainties in the current atmospheric budgets of these gases. Through synthesizing relevant studies, we propose that aggregates, as relatively separate micro-environments embedded in a complex soil matrix, can be viewed as biogeochemical reactors of GHGs. Aggregate reactivity is determined by both aggregate size (which determines the reactor size) and the bulk soil environment including both biotic and abiotic factors (which further influence the reaction conditions). With a systematic, dynamic view of the soil system, implications of aggregate reactors for soil-atmosphere GHG exchange are determined by both an individual reactor's reactivity and dynamics in aggregate size distributions. Emerging evidence supports the contention that aggregate reactors significantly influence soil-atmosphere GHG exchange and may have global implications for carbon and nitrogen cycling. In the context of increasingly frequent and severe disturbances, we advocate more analyses of GHG activities at the aggregate scale. To complement data on aggregate reactors, we suggest developing bottom-up aggregate-based models (ABMs) that apply a trait-based approach and incorporate soil system heterogeneity.

Root morphology and mycorrhizal symbioses together shape nutrient foraging strategies of temperate trees.

Photosynthesis by leaves and acquisition of water and minerals by roots are required for plant growth, which is a key component of many ecosystem functions. Although the role of leaf functional traits in photosynthesis is generally well understood, the relationship of root functional traits to nutrient uptake is not. In particular, predictions of nutrient acquisition strategies from specific root traits are often vague. Roots of nearly all plants cooperate with mycorrhizal fungi in nutrient acquisition. Most tree species form symbioses with either arbuscular mycorrhizal (AM) or ectomycorrhizal (EM) fungi. Nutrients are distributed heterogeneously in the soil, and nutrient-rich "hotspots" can be a key source for plants. Thus, predicting the foraging strategies that enable mycorrhizal root systems to exploit these hotspots can be critical to the understanding of plant nutrition and ecosystem carbon and nutrient cycling. Here, we show that in 13 sympatric temperate tree species, when nutrient availability is patchy, thinner root species alter their foraging to exploit patches, whereas thicker root species do not. Moreover, there appear to be two distinct pathways by which thinner root tree species enhance foraging in nutrient-rich patches: AM trees produce more roots, whereas EM trees produce more mycorrhizal fungal hyphae. Our results indicate that strategies of nutrient foraging are complementary among tree species with contrasting mycorrhiza types and root morphologies, and that predictable relationships between below-ground traits and nutrient acquisition emerge only when both roots and mycorrhizal fungi are considered together.

Environmental Controls on Soil Microbial Communities in a Seasonally Dry Tropical Forest.

Several studies have shown that rainfall seasonality, soil heterogeneity, and increased nitrogen (N) deposition may have important effects on tropical forest function. However, the effects of these environmental controls on soil microbial communities in seasonally dry tropical forests are poorly understood. In a seasonally dry tropical forest in the Yucatan Peninsula (Mexico), we investigated the influence of soil heterogeneity (which results in two different soil types, black and red soils), rainfall seasonality (in two successive seasons, wet and dry), and 3 years of repeated N enrichment on soil chemical and microbiological properties, including bacterial gene content and community structure. The soil properties varied with the soil type and the sampling season but did not respond to N enrichment. Greater organic matter content in the black soils was associated with higher microbial biomass, enzyme activities, and abundances of genes related to nitrification (amoA) and denitrification (nirK and nirS) than were observed in the red soils. Rainfall seasonality was also associated with changes in soil microbial biomass and activity levels and N gene abundances. Actinobacteria, Proteobacteria, Firmicutes, and Acidobacteria were the most abundant phyla. Differences in bacterial community composition were associated with soil type and season and were primarily detected at higher taxonomic resolution, where specific taxa drive the separation of communities between soils. We observed that soil heterogeneity and rainfall seasonality were the main correlates of soil bacterial community structure and function in this tropical forest, likely acting through their effects on soil attributes, especially those related to soil organic matter and moisture content.IMPORTANCE Understanding the response of soil microbial communities to environmental factors is important for predicting the contribution of forest ecosystems to global environmental change. Seasonally dry tropical forests are characterized by receiving less than 1,800 mm of rain per year in alternating wet and dry seasons and by high heterogeneity in plant diversity and soil chemistry. For these reasons, N deposition may affect their soils differently than those in humid tropical forests. This study documents the influence of rainfall seasonality, soil heterogeneity, and N deposition on soil chemical and microbiological properties in a seasonally dry tropical forest. Our findings suggest that soil heterogeneity and rainfall seasonality are likely the main factors controlling soil bacterial community structure and function in this tropical forest. Nitrogen enrichment was likely too low to induce significant short-term effects on soil properties, because this tropical forest is not N limited.

The effectiveness of cutoff walls to control saltwater intrusion in multi-layered coastal aquifers: Experimental and numerical study.

The objective of this study was to examine the performance of cutoff walls in controlling saltwater intrusion in stratified heterogeneous coastal aquifers. Numerical and laboratory experiments were completed in laboratory-scale aquifer where the effectiveness of cutoff walls was assessed in three different configurations, including a homogeneous scenario, a stratified aquifer with high K-low K-high K pattern (case HLH) and another stratified aquifer with low K-high K-low K pattern (case LHL). The results show that the cutoff wall was effective in reducing the saltwater wedge in all the investigated cases of layered-aquifers with toe length reduction of up to 43%. The wall exhibited more wedge reduction in shallower than steeper hydraulic gradients. However, the soil stratification appeared to lessen the overall performance of the wall compared to the homogeneous case. The aquifer stratification disrupted the flow dynamics, and thus affected the freshwater velocity at the wall opening to various degrees, depending on the layering pattern. The presence of an interlayer of low k (case HLH) inhibited the downward movement of the freshwater towards the wall opening, and thus decreasing the repulsion ability of the wall. Moreover, the presence of an underlying low permeability layer (case LHL) was found to obstruct the freshwater flow in the lower part of the aquifer, thereby slowing down the velocity through the wall opening. Numerical analysis of other layering patterns of monotonically increasing/decreasing permeability from top to bottom showed that the cutoff wall remained effective in repulsing the seawater wedge.

Spatial heterogeneity of plant-soil feedback affects root interactions and interspecific competition.

Plant-soil feedback is receiving increasing interest as a factor influencing plant competition and species coexistence in grasslands. However, we do not know how spatial distribution of plant-soil feedback affects plant below-ground interactions. We investigated the way in which spatial heterogeneity of soil biota affects competitive interactions in grassland plant species. We performed a pairwise competition experiment combined with heterogeneous distribution of soil biota using four grassland plant species and their soil biota. Patches were applied as quadrants of 'own' and 'foreign' soils from all plant species in all pairwise combinations. To evaluate interspecific root responses, species-specific root biomass was quantified using real-time PCR. All plant species suffered negative soil feedback, but strength was species-specific, reflected by a decrease in root growth in own compared with foreign soil. Reduction in root growth in own patches by the superior plant competitor provided opportunities for inferior competitors to increase root biomass in these patches. These patterns did not cascade into above-ground effects during our experiment. We show that root distributions can be determined by spatial heterogeneity of soil biota, affecting plant below-ground competitive interactions. Thus, spatial heterogeneity of soil biota may contribute to plant species coexistence in species-rich grasslands.

Contrasts between whole-plant and local nutrient levels determine root growth and death in Ailanthus altissima (Simaroubaceae).

UNLABELLED: • PREMISE OF THE STUDY: There is an ongoing debate about the importance of whole-plant control vs. local modular mechanisms for root growth. We conducted a split-root experiment with different patch/background levels of nitrogen to examine whether local root growth and death are controlled by local resource levels or at the whole-plant level.• METHODS: Three microrhizotrons with 0, 10, and 100 µg N/g growth medium levels (74 g growth medium each) were attached to pots of high or low soil N in which one Ailanthus altissima individual was growing. One fine root was guided into each of the microrhizotrons and photographed every 4 d. Plants were harvested after 28 d; root growth and mortality in the microrhizotrons were recorded. Changes in root length, number of laterals, and interlateral length were determined from the photos and analyzed.• KEY RESULTS: While overall plant growth was influenced by background N level, both patch and background N levels influenced root growth and mortality in patches. Local roots proliferated most when the patch N level was high and background level low, and they proliferated least and showed highest mortality when patch N was low and the background level high.• CONCLUSIONS: The fate of roots growing in a patch is influenced by the resource environment of the plant's other roots as well as the resource levels in the patch itself. Thus, the growth and death of roots in patches is determined by both modular and whole-plant mechanisms.

Root plasticity versus elasticity - when are responses acclimative?

Spatiotemporal soil heterogeneity and the resulting edaphic stress cycles can be decisive for crop growth. However, our understanding of the acclimative value of root responses to heterogeneous soil conditions remains limited. We outline a framework to evaluate the acclimative value of root responses that distinguishes between stress responses that are persistent and reversible upon stress release, termed 'plasticity' and 'elasticity', respectively. Using energy balances, we provide theoretical evidence that the advantage of plasticity over elasticity increases with the number of edaphic stress cycles and if responses lead to comparatively high energy gains. Our framework provides a conceptual basis for assessing the acclimative value of root responses to soil heterogeneity and can catalyse research on crop adaptations to heterogeneous belowground environments.