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1.

Background

In order to maintain high yields while saving water and preserving non-renewable resources and thus limiting the use of chemical fertilizer, it is crucial to select plants with more efficient root systems. This could be achieved through an optimization of both root architecture and root uptake ability and/or through the improvement of positive plant interactions with microorganisms in the rhizosphere. The development of devices suitable for high-throughput phenotyping of root structures remains a major bottleneck.

Results

Rhizotrons suitable for plant growth in controlled conditions and non-invasive image acquisition of plant shoot and root systems (RhizoTubes) are described. These RhizoTubes allow growing one to six plants simultaneously, having a maximum height of 1.1 m, up to 8 weeks, depending on plant species. Both shoot and root compartment can be imaged automatically and non-destructively throughout the experiment thanks to an imaging cabin (RhizoCab). RhizoCab contains robots and imaging equipment for obtaining high-resolution pictures of plant roots. Using this versatile experimental setup, we illustrate how some morphometric root traits can be determined for various species including model (Medicago truncatula), crops (Pisum sativum, Brassica napus, Vitis vinifera, Triticum aestivum) and weed (Vulpia myuros) species grown under non-limiting conditions or submitted to various abiotic and biotic constraints. The measurement of the root phenotypic traits using this system was compared to that obtained using “classic” growth conditions in pots.

Conclusions

This integrated system, to include 1200 Rhizotubes, will allow high-throughput phenotyping of plant shoots and roots under various abiotic and biotic environmental conditions. Our system allows an easy visualization or extraction of roots and measurement of root traits for high-throughput or kinetic analyses. The utility of this system for studying root system architecture will greatly facilitate the identification of genetic and environmental determinants of key root traits involved in crop responses to stresses, including interactions with soil microorganisms.
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2.

Context

Animals selectively use landscapes to meet their energetic needs, and trade-offs in habitat use may depend on availability and environmental conditions. For example, habitat selection at high temperatures may favor thermal cover at the cost of reduced foraging efficiency under consistently warm conditions.

Objective

Our objective was to examine habitat selection and space use in distinct populations of moose (Alces alces). Hypothesizing that endotherm fitness is constrained by heat dissipation efficiency, we predicted that southerly populations would exhibit greater selection for thermal cover and reduced selection for foraging habitat.

Methods

We estimated individual step selection functions with shrinkage for 134 adult female moose in Minnesota, USA, and 64 in Ontario, Canada, to assess habitat selection with variation in temperature, time of day, and habitat availability. We averaged model coefficients within each site to quantify selection strength for habitats differing in forage availability and thermal cover.

Results

Moose in Ontario favored deciduous and mixedwood forest, indicating selection for foraging habitat across both diel and temperature. Habitat selection patterns of moose in Minnesota were more dynamic and indicated time- and temperature-dependent trade-offs between use of foraging habitat and thermal cover.

Conclusions

We detected a scale-dependent functional response in habitat selection driven by the trade-off between selection for foraging habitat and thermal cover. Landscape composition and internal state interact to produce complex patterns of space use, and animals exposed to increasingly high temperatures may mitigate fitness losses from reduced foraging efficiency by increasing selection for foraging habitat in sub-prime foraging landscapes.
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3.

Context

Competitive interactions potentially play an important role in structuring bird communities. It is unclear how differences in functional traits influence the niche dimensions of highly mobile waterbird species, particularly when they co-exist in spatiotemporally heterogeneous communities.

Objectives

We investigated the inter-relationships between waterbird trait groupings (movement, dietary and foraging habitat) and environmental variable groupings (rainfall, land cover, vegetation structure and water quality). Specifically, we tested whether the scale of environmental variables filtered movement traits and whether these traits operated in conjunction with dietary and foraging habitat traits to form distinct ecological niches in waterbirds.

Methods

We conducted waterbird and environmental variable surveys in 60 sites, sampled seven times each at bimonthly intervals, in KwaZulu-Natal, South Africa. Trait-environment relationships were tested using a combination of RLQ and fourth-corner analyses.

Results

Several significant trait-environment relationships emerged in bivariate correlations and multivariate ordination space. Movement traits correlated with the scale of environmental variables; migrant and nomadic species responded to broad scale environmental variables. Vegetation structure and land cover were particularly important in explaining the abundance of species foraging in emergent vegetation. Three groups emerged along a gradient in multivariate ordination space providing evidence for ecological niche separation of waterbirds with different movement traits.

Conclusions

Our findings suggest that the scale of landscape resources can act as a filter of movement traits, and that in conjunction with dietary and foraging traits, waterbirds with different movement traits occupy distinct ecological niches.
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4.

Context

The conversion of natural environments into agricultural land has profound effects on the composition of the landscape, often resulting in a mosaic of human-altered and natural habitats. The response to these changes may however vary among organisms. Bats are highly vagile, and their requirements often imply the use of distinct habitats, which they select responding to both landscape and local features.

Objectives

We aimed to identify which features influence bat richness and activity within Baixo Vouga Lagunar, a heterogeneous landscape located on the Central-North Portuguese coast, and to investigate if that influence varies across a gradient of focal scales.

Methods

We sampled bats acoustically, while simultaneously sampling insects with light traps. We assessed the relationships between species richness, bat activity, and activity of eco-morphological guilds with landscape and local features, across four scales.

Results

Our results revealed both scale- and guild-dependent responses of bats to landscape and local features. At broader scales we found positive associations between open-space foraging bats and habitat heterogeneity and between edge-space foraging bats and greater edge lengths. Woodland cover and water availability at an intermediate scale and weather conditions and insect abundance at a local scale were the factors that mostly influenced the response variables.

Conclusions

Globally, our results suggest that bats are sensitive to local resource availability and distribution, while simultaneously reacting to landscape features acting at coarser scales. Finally, our results suggest that the responses given by bats are guild-dependent, and some habitats act as keystone structures for bats within this mosaic.
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5.

Background

Genetic studies on the molecular mechanisms of the regulation of root growth require the characterisation of a specific root phenotype to be linked with a certain genotype. Such studies using classical labour-intensive methods are severely hindered due to the technical limitations that are associated with the impeded observation of the root system of a plant during its growth. The aim of the research presented here was to develop a reliable, cost-effective method for the analysis of a plant root phenotype that would enable the precise characterisation of the root system architecture of cereals.

Results

The presented method describes a complete system for automatic supplementation and continuous sensing of culture solution supplied to plants that are grown in transparent tubes containing a solid substrate. The presented system comprises the comprehensive pipeline consisting of a modular-based and remotely-controlled plant growth system and customized imaging setup for root and shoot phenotyping. The system enables an easy extension of the experimental capacity in order to form a combined platform that is comprised of parallel modules, each holding up to 48 plants. The conducted experiments focused on the selection of the most suitable conditions for phenotyping studies in barley: an optimal size of the glass beads, diameters of the acrylic tubes, composition of a medium, and a rate of the medium flow.

Conclusions

The developed system enables an efficient, accurate and highly repeatable analysis of the morphological features of the root system of cereals. Because a simple and fully-automated control system is used, the experimental conditions can easily be normalised for different species of cereals. The scalability of the module-based system allows its capacity to be adjusted in order to meet the requirements of a particular experiment.
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6.

Context

Many arboreal mammals in Neotropical forests are important seed dispersers that influence the spatial patterns of tree regeneration via their movement patterns, which in turn are determined by the canopy structure of the forest itself. However, the relationship between arboreal mammal movement and canopy structure is poorly understood, due in large part to the complexity of quantifying arboreal habitat structure.

Objectives

We relate detailed movement trajectories of three sympatric primate species to attributes of canopy structure derived from airborne light detection and ranging (LiDAR) in order to understand the role of structure in arboreal movement in the tropical moist forest of Barro Colorado Island, Panama.

Methods

We used high-resolution LiDAR to quantify three-dimensional attributes of the forest canopy of the entire island, high-resolution GPS tracking to map the movement patterns of the monkey species, and step selection functions to relate movement decisions to canopy attributes.

Results

We found that movement decisions were correlated with canopy height and distance to gaps, which indicate forest maturity and lateral connectivity, in all three species. In the two faster-moving species, step selection was also correlated with the thickness of the crown layer and the density of vegetation within the crown.

Conclusions

The correlations detected are fully in line with known differences in the locomotor adaptations and movement strategies of the study species, and directly reflect maximization of energetic efficiency and ability to escape from predators. Quantification of step selection in relation to structure thus provides insight into the ways in which arboreal animals use their environment.
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7.

Background

Plant roots release a variety of organic compounds into the soil which alter the physical, chemical and biological properties of the rhizosphere. Root exudates are technically challenging to measure in soil because roots are difficult to access and exudates can be bound by minerals or consumed by microorganisms. Exudates are easier to measure with hydroponically-grown plants but, even here, simple compounds such as sugars and organic acids can be rapidly assimilated by microorganisms. Sterile hydroponic systems avoid this shortcoming but it is very difficult to maintain sterility for long periods especially for larger crop species. As a consequence, studies often use small model species such as Arabidopsis to measure exudates or use seedlings of crop plants which only have immature roots systems.

Results

We developed a simple hydroponic system for cultivating large crop plants in sterile conditions for more than 30 days. Using this system wheat (Triticum aestivum L.) and barley (Hordeum vulgare L.) plants were grown in sterile conditions for 30 days by which time they had reached the six-leaf stage and developed mature root systems with seminal, nodal and lateral roots. To demonstrate the utility of this system we characterized the aluminium-activated exudation of malate from the major types of wheat roots for the first time. We found that all root types measured released malate but the amounts were two-fold greater from the seminal and nodal axile roots compared with the lateral roots. Additionally, we showed that this sterile growth system could be used to collect exudates from intact whole root systems of barley.

Conclusions

We developed a simple hydroponic system that enables cereal plants to be grown in sterile conditions for longer periods than previously recorded. Using this system we measured, for the first time, the aluminium-activated efflux of malate from the major types of wheat roots. We showed the system can also be used for collecting exudates from intact root systems of 30-day-old barley plants. This hydroponic system can be modified for various purposes. Importantly it enables the study of exudates from crop species with mature root systems.
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8.

Context

An increasing number of studies have investigated the impact of environmental heterogeneity on faunal assemblages when measured at multiple spatial scales. Few studies, however, have considered how the effects of heterogeneity on fauna vary with the spatial scale at which the response variable is characterised.

Objectives

We investigated the relationship between landscape properties in a region characterised by diverse fire mosaics, and the structure and composition of avian assemblages measured at both the site- (1 ha) and landscape-scale (100 ha).

Methods

We surveyed birds and calculated spatial landscape properties in sub-tropical woodlands of central Queensland, Australia.

Results

Environmental heterogeneity, as measured by topographic complexity, was consistently important for bird species richness and composition. However, the explanatory power of topographic complexity varied depending on the spatial scale and the component of diversity under investigation. We found different correlates of richness within particular foraging guilds depending on the scale at which richness was measured. Extent of long-unburnt habitat (>10 years since fire) was the most important variable for the landscape-scale richness of frugivores, insectivores and canopy feeders, whereas environmental heterogeneity in the surrounding landscape was more important for site-scale richness of these foraging guilds.

Conclusions

The response of species richness to landscape characteristics varies among scales, and among components of diversity. Thus, depending on the scale at which a biodiversity conservation goal is conceptualised—maximising richness at a site, or across a landscape—different landscape management approaches may be preferred.
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9.

Context

Disturbances create spatial variation in environments that may influence animal foraging. Granivory by rodents can influence seed supply and thus plant establishment. However, effects of disturbance patterns on rodent seed removal in western North American conifer forests are generally unknown.

Objectives

We conducted a study in lodgepole pine (Pinus contorta var. latifolia) forests of Greater Yellowstone (Wyoming, USA) to answer: (1) How do seed removal and rodent activity vary between recently burned and adjacent unburned forests and with distance from fire perimeter? (2) Which microhabitat conditions explain variability in seed removal and rodent activity?

Methods

One or two years after wildfires, we established transects (n = 23) with four stations each: at 10 and 40 m from the fire perimeter in both burned and unburned forest. At stations, we deployed trays with lodgepole pine seeds and cameras pointed at trays for 28 days and quantified habitat structure and seed abundance.

Results

Seed removal, which averaged 85%, and diurnal rodent activity did not differ between burned and unburned forests or with distance from the fire perimeter; however, nocturnal rodent activity was lower in burned forests. Seed removal and diurnal rodent activity were not associated with any microhabitat conditions we measured. However, nocturnal rodent activity was associated with microhabitat in both burned and unburned forests.

Conclusions

High seed removal rates suggested that rodent foraging was not reduced by high-severity wildfire. If observed seed removal represents natural conditions, post-dispersal seed predation could influence post-fire recruitment of a widespread foundation tree species.
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10.
11.

Context

Despite decades of research, there is an intense debate about the consistency of the hump-shaped pattern describing the relationship between diversity and disturbance as predicted by the intermediate disturbance hypothesis (IDH). Previous meta-analyses have not explicitly considered interactive effects of disturbance frequency and intensity of disturbance on plant species diversity in terrestrial landscapes.

Objective

We conducted meta-analyses to test the applicability of IDH by simultaneously examining the relationship between species richness, disturbance frequency (quantified as time since last disturbance as originally proposed) and intensity of disturbance in forest landscapes.

Methods

The effects of disturbance frequency, intensity, and their interaction on species richness was evaluated using a mixed-effects model.

Results

We found that species richness peaks at intermediate frequency after both high and intermediate disturbance intensities, but the richness-frequency relationship differed between intensity classes.

Conclusions

Our study highlights the need to measure multiple disturbance components that could help reconcile conflicting empirical results on the effect of disturbance on plant species diversity.
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12.

Context

The spatial distribution of non-substitutable resources implies diverging predictions for animal movement patterns. At broad scales, animals should respond to landscape complementation by selecting areas where resource patches are close-by to minimize movement costs. Yet at fine scales, central place effects lead to the depletion of patches that are close to one another and that should ultimately be avoided by consumers.

Objectives

We developed a multi-scale resource selection framework to test whether animal movement is driven by landscape complementation or resource depletion and identify at which spatial scale these processes are relevant from an animal’s perspective.

Methods

During the dry season, surface water and forage are non-substitutable resources for African elephants. Eight family herds were tracked using GPS loggers in Hwange National Park, Zimbabwe. We explained habitat selection during foraging trips by mapping surface water at two scales with gaussian kernels of varying widths placed over each waterhole.

Results

Unexpectedly, elephants select areas with low waterhole density at both fine scales (< 1 km) and broad scales (5–7 km). Selection is stronger when elephants forage far away from water, even more so as the dry season progresses.

Conclusions

Elephant selection of low waterhole density areas suggests that resource depletion around multiple central places is the main driver of their habitat selection. By identifying the scale at which animals respond to waterhole distribution we provide a template for water management in arid and semi-arid landscapes that can be tailored to match the requirements and mobility of free ranging wild or domestic species.
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13.

Context

Forest insect outbreaks are influenced by ecological processes operating at multiple spatial scales, including host-insect interactions within stands and across landscapes that are modified by regional-scale variations in climate. These drivers of outbreak dynamics are not well understood for the western spruce budworm, a defoliator that is native to forests of western North America.

Objectives

Our aim was to assess how processes across multiple spatial scales drive western spruce budworm outbreak dynamics. Our objective was to assess the relative importance and influence of a set of factors covering the stand, landscape, and regional scales for explaining spatiotemporal outbreak patterns in British Columbia, Canada.

Methods

We used generalized linear mixed effect models within a multi-model interference framework to relate annual budworm infestation mapped from Landsat time series (1996–2012) to sets of stand-, landscape-, and regional-scale factors derived from forest inventory data, GIS analyses, and climate models.

Results

Outbreak patterns were explained well by our model (R 2 = 93%). The most important predictors of infestation probability were the proximity to infestations in the previous year, landscape-scale host abundance, and dry autumn conditions. While stand characteristics were overall less important predictors, we did find infestations were more likely amongst pure Douglas-fir stands with low site indices and high crown closure.

Conclusions

Our findings add to growing empirical evidence that insect outbreak dynamics are driven by multi-scaled processes. Forest management planning to mitigate the impacts of budworm outbreaks should thus consider landscape- and regional-scale factors in addition to stand-scale factors.
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14.

Context

Landscape heterogeneity (the composition and configuration of different landcover types) plays a key role in shaping woodland bird assemblages in wooded-agricultural mosaics. Understanding how species respond to landscape factors could contribute to preventing further decline of woodland bird populations.

Objective

To investigate how woodland birds with different species traits respond to landscape heterogeneity, and to identify whether specific landcover types are important for maintaining diverse populations in wooded-agricultural environments.

Methods

Birds were sampled from woodlands in 58 2 × 2 km tetrads across southern Britain. Landscape heterogeneity was quantified for each tetrad. Bird assemblage response was determined using redundancy analysis combined with variation partitioning and response trait analyses.

Results

For woodland bird assemblages, the independent explanatory importance of landscape composition and landscape configuration variables were closely interrelated. When considered simultaneously during variation partitioning, the community response was better represented by compositional variables. Different species responded to different landscape features and this could be explained by traits relating to woodland association, foraging strata and nest location. Ubiquitous, generalist species, many of which were hole-nesters or ground foragers, correlated positively with urban landcover while specialists of broadleaved woodland avoided landscapes containing urban areas. Species typical of coniferous woodland correlated with large conifer plantations.

Conclusions

At the 2 × 2 km scale, there was evidence that the availability of resources provided by proximate landcover types was highly important for shaping woodland bird assemblages. Further research to disentangle the effects of composition and configuration at different spatial scales is advocated.
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15.

Context

In the Rio Grande Plains of southern Texas, subtropical savanna vegetation is characterized by a two-phase pattern consisting of discrete woody patches embedded within a C4 grassland matrix. Prior trench transect studies have suggested that, on upland portions of the landscape, large woody patches (groves) occur on non-argillic inclusions, while small woody patches (clusters) are dispersed among herbaceous vegetation where the argillic horizon is present.

Objective

To test whether spatial heterogeneity of subsurface soil texture drives the landscape-scale pattern of woody patches in this subtropical savanna.

Methods

Landscape-scale spatial patterns of soil texture were quantified by taking spatially-specific soil samples to a depth of 1.2 m in a 160 m × 100 m plot. Kriged maps of soil texture were developed, and the locations of non-argillic inclusions were mapped.

Results

Visual comparison of kriged maps of soil texture to a high resolution aerial photograph of the study area revealed that groves were present exclusively where the non-argillic inclusions were present. This clear visual relationship was further supported by positive correlations between soil sand concentration in the lower soil layers and total fine root biomass which mapped the locations of groves.

Conclusions

Subsurface non-argillic inclusions may favor the establishment and persistence of groves by enabling root penetration deeper into the profile, providing greater access to water and nutrients that are less accessible on those portions of the landscape where the argillic horizon is present, thereby regulating the distribution of grove vegetation and structuring the evolution of this landscape.
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16.

Context

Landscape resistance is vital to connectivity modeling and frequently derived from resource selection functions (RSFs). RSFs estimate relative probability of use and tend to focus on understanding habitat preferences during slow, routine animal movements (e.g., foraging). Dispersal and migration, however, can produce rarer, faster movements, in which case models of movement speed rather than resource selection may be more realistic for identifying habitats that facilitate connectivity.

Objective

To compare two connectivity modeling approaches applied to resistance estimated from models of movement rate and resource selection.

Methods

Using movement data from migrating elk, we evaluated continuous time Markov chain (CTMC) and movement-based RSF models (i.e., step selection functions [SSFs]). We applied circuit theory and shortest random path (SRP) algorithms to CTMC, SSF and null (i.e., flat) resistance surfaces to predict corridors between elk seasonal ranges. We evaluated prediction accuracy by comparing model predictions to empirical elk movements.

Results

All connectivity models predicted elk movements well, but models applied to CTMC resistance were more accurate than models applied to SSF and null resistance. Circuit theory models were more accurate on average than SRP models.

Conclusions

CTMC can be more realistic than SSFs for estimating resistance for fast movements, though SSFs may demonstrate some predictive ability when animals also move slowly through corridors (e.g., stopover use during migration). High null model accuracy suggests seasonal range data may also be critical for predicting direct migration routes. For animals that migrate or disperse across large landscapes, we recommend incorporating CTMC into the connectivity modeling toolkit.
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17.

Context

In Europe, policy measures are starting to emerge that promote multifunctional farming systems and delivery of ecosystem services besides food production. Effectiveness of these policy instruments have to deal with ecological, economic and social complexities and with complexities in individual decisions of local actors leading to system shifts.

Objective

The objective of this paper is to discover the most important social and/or economic drivers that cause farm systems to shift between a monofunctional (providing food) and a multifunctional state (providing food and natural pest regulation).

Methods

Using a cellular automata model, we simulated decisions of individual farmers to shift between a mono-and multifunctional state through time, based on their behaviour type and on financial and social consequences. Collaboration of multifunctional farmers at a landscape scale is a precondition to provide a reliable level of natural pest regulation.

Results

Costs of applying green infrastructure was an important driver for the size and the conversion rate of shifts between mono-and multifunctional farming systems. Shifts towards multifunctional farming were enhanced by a higher motivation of farmers to produce sustainably, while shifts (back) to a monofunctional state was enhanced by a low social cohesion between multifunctional farmers.

Conclusions

These results suggest that in order to develop a multifunctional farming system, individual farmers should act counterintuitively to their conventional farming environment. To maintain a multifunctional farming system, social cohesion between multifunctional farmers is most relevant. Financial aspects are important in both shifts.
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18.

Context

Quantitative models of forest dynamics have followed a progression toward methods with increased detail, complexity, and spatial extent.

Objectives

We highlight milestones in the development of forest dynamics models and identify future research and application opportunities.

Methods

We reviewed milestones in the evolution of forest dynamics models from the 1930s to the present with emphasis on forest growth and yield models and forest landscape models We combined past trends with emerging issues to identify future needs.

Results

Historically, capacity to model forest dynamics at tree, stand, and landscape scales was constrained by available data for model calibration and validation; computing capacity; model applicability to real-world problems; and ability to integrate biological, social, and economic drivers of change. As computing and data resources improved, a new class of spatially explicit forest landscape models emerged.

Conclusions

We are at a point of great opportunity in development and application of forest dynamics models. Past limitations in computing capacity and in data suitable for model calibration or evaluation are becoming less restrictive. Forest landscape models, in particular, are ready to transition to a central role supporting forest management, planning, and policy decisions.

Recommendations

Transitioning forest landscape models to a central role in applied decision making will require greater attention to evaluating performance; building application support staffs; expanding the included drivers of change, and incorporating metrics for social and economic inputs and outputs.
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19.

Context

In response to predominantly local and private approaches to landscape change, landscape ecologists should critically assess the multiscalar influences on landscape design.

Objectives

This study develops a governance framework for Nassauer and Opdam’s “Design-in-Science” model. Its objective is to create an approach for examining hierarchical constraints on landscape design in order to investigate linkages among urban greening initiatives, patterns of landscape change, and the broader societal values driving those changes. It aims to provide an integrative and actionable approach for landscape sustainability science.

Methods

This framework is examined through an ethnographic study of public policy processes surrounding the urban tree initiatives in Boston, MA; Philadelphia, PA; and Baltimore, MD.

Results

These initiatives demonstrate the impact of political and economic decentralization on urban landscape patterns. Their collaborative governance approach incorporates diverse resources to implement programming at a fine-scale. The predominant tree giveaway program fragments the urban and regional forest.

Conclusion

Spatial and temporal fragmentation undermines the long-term security of urban greening programs, and it suggests reconsideration of the role of state regimes in driving broad scale spatial planning.
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20.

Background

Root systems are highly plastic and adapt according to their soil environment. Studying the particular influence of soils on root development necessitates the adaptation and evaluation of imaging methods for multiple substrates. Non-invasive 3D root images in soil can be obtained using magnetic resonance imaging (MRI). Not all substrates, however, are suitable for MRI. Using barley as a model plant we investigated the achievable image quality and the suitability for root phenotyping of six commercially available natural soil substrates of commonly occurring soil textures. The results are compared with two artificially composed substrates previously documented for MRI root imaging.

Results

In five out of the eight tested substrates, barley lateral roots with diameters below 300 µm could still be resolved. In two other soils, only the thicker barley seminal roots were detectable. For these two substrates the minimal detectable root diameter was between 400 and 500 µm. Only one soil did not allow imaging of the roots with MRI. In the artificially composed substrates, soil moisture above 70% of the maximal water holding capacity (WHCmax) impeded root imaging. For the natural soil substrates, soil moisture had no effect on MRI root image quality in the investigated range of 50–80% WHCmax.

Conclusions

Almost all tested natural soil substrates allowed for root imaging using MRI. Half of these substrates resulted in root images comparable to our current lab standard substrate, allowing root detection down to a diameter of 300 µm. These soils were used as supplied by the vendor and, in particular, removal of ferromagnetic particles was not necessary. With the characterization of different soils, investigations such as trait stability across substrates are now possible using noninvasive MRI.
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