No abstract

Contradictory evidence exists regarding whether and to which extend roots change soil structure in their vicinity. Here we attempt to reconcile disparate views allowing for the two-way interaction between soil structure and root traits, i.e. changes in soil structure due to plants and changes in root growth due to soil structure. Porosity gradients extending from the root/biopore surface into the bulk soil were investigated with X-ray µCT for undisturbed soil samples from a field chronosequence as well as for a laboratory experiment with Zea mays growing into three different bulk densities. An image analysis protocol was developed, which enabled a fast analysis of the large sample pool (n > 300) at a resolution of 19 µm. Lab experiment showed that growing roots only compact the surrounding soil if macroporosity is low and dominated by isolated pores. When roots can grow into a highly connected macropore system showing high connectivity the rhizosphere is more porous compared to the bulk soil. A compaction around roots/biopores in the field chronosequence was only observed in combination with high root/biopore length densities. We conclude that roots compact the rhizosphere only if the initial soil structure does not offer a sufficient volume of well-connected macropores.

Connectivity is one of the most important parameters to quantify pore structure and link it to soil functions. One of the great challenges in quantifying connectivity with X‐ray microtomography (X‐ray μCT) is that high resolution, as required for small pores, can only be achieved in small samples in which the connectivity of larger pores can no longer be quantified in a meaningful way. The objective of this study was to investigate the changes in pore connectivity with changing sample size, covering a range of analysed pore diameters of more than three orders of magnitude. With this approach, we wanted to address whether pore types formed by different processes in an agricultural chronosequence leave characteristic traces in certain connectivity metrics. The Euler number, χ, and the connection probability of two random points within the pore system, that is, the Γ‐indicator, were determined as a function of minimum pore diameter. The results show that characteristic signatures of certain pore types overlap with scale artifacts in the connectivity functions. The Γ‐indicator, gives highly biased information in small samples. Therefore, we developed a new method for a joint‐Γ‐curve that merges information from three samples sizes. However, χ does not require such a scale fusion. It can be used to define characteristic size ranges for pore types and is very sensitive to the occurrence of bottle necks. Our findings suggest a joint evaluation of both connectivity metrics to disentangle different pore types with χ and to identify the contribution of different pore types to the overall pore connectivity with Γ. This evaluation on the chronosequence showed that biopores mainly connect pores of diameters between 0.5 and 0.1 mm. This was not coupled with an increase in pore volume. In contrast, tillage led to a shift of pores of diameter >0.05 mm towards pores of diameter >0.20 mm and thus increased connectivity of pores >0.20 mm. This work underlines the importance of accounting for the scale dependence of connectivity measures and provides a methodological approach for doing so. Highlights Scale dependence of connectivity metrics needs to be accounted for. Connectivity metrics can be used to disentangle different pore types across scales. Roots mainly connect the pore system between 0.1 and 0.5 mm. A joint Γ‐connectivity function can be constructed that is free of scale artifacts.

No abstract

Soil is a heterogeneous mixture of various organic and inorganic parent materials. Major soil functions are driven by their quality, quantity and spatial arrangement, resulting in soil structure. Physical protection of organic matter (OM) in this soil structure is considered as a vital mechanism for stabilizing organic carbon turnover, an important soil function in times of climate change. Herein, we present a technique for the correlative analysis of 2D imaging visible light near-infrared spectroscopy and 3D X-ray computed microtomography (µCT) to investigate the interplay of biogeochemical properties and soil structure in undisturbed soil samples. Samples from the same substrate but different soil management and depth (no-tilled topsoil, tilled topsoil and subsoil) were compared in order to evaluate this method in a diversely structured soil. Imaging spectroscopy is generally used to qualitatively and quantitatively identify OM with high spatial resolution, whereas 3D X-ray µCT provides high-resolution information on pore characteristics. The unique combination of these techniques revealed that, in undisturbed samples, OM can be found mainly at greater distances from macropores and close to biopores. However, alterations were observed because of disturbances by tillage. The correlative application of imaging infrared spectroscopic and X-ray µCT analysis provided new insights into the biochemical processes affected by soil structural changes.

Apple replant disease (ARD) is the phenomenon of soil decline occurring after repeated planting of apple trees at the same site. This study aimed to elucidate whether ARD is systemic, i.e. whether the contact of parts of the root system with ARD soil causes the whole plant to show poor shoot and root growth. A split-root experiment was conducted with seedlings of ‘M26’, offering the same plant for its root system the choice between the substrates ARD soil (+ARD), γ-sterilized ARD soil (-ARD) or soil from a grass parcel (Control) with the following combinations: +ARD/+ARD, -ARD/-ARD; +ARD/-ARD; +ARD/Control. Root growth was analysed throughout the 34-day growing period. Samples from bulk, rhizosphere and rhizoplane soil were collected separately for each compartment, and analysed by fingerprints of 16S rRNA gene or ITS fragments amplified from total community (TC) DNA. The response of the plant to +ARD was not systemic as root growth in -ARD compartment was always superior to root growth in +ARD soil. Crosswise 15N-labelling of the N-fertilizer applied to the split-root compartments showed that nitrate-N uptake efficiency was higher for roots in -ARD soil compared to those in +ARD. Bacterial and fungal community composition in the rhizoplane and rhizosphere of the same plants differed significantly between the compartments containing +ARD/-ARD or +ARD/Control. The strongest differences between the bacterial fingerprints were observed in the rhizoplane and rhizosphere. Bacterial genera with increased abundance in response to ARD were mainly Streptomyces but also Sphingobium, Novosphingobium, Rhizobium, Lysobacter and Variovorax. The strongest differences between the fungal fingerprints were observed in bulk soil. Our data showed that the response of the apple plant to ARD soil is local and not systemic.

Apple replant disease (ARD) occurs worldwide in apple orchards and nurseries and leads to a severe growth and productivity decline. Despite research on the topic, its causality remains unclear. In a split-root experiment, we grew ARD-susceptible ‘M26’ apple rootstocks in different substrates combinations (+ARD: apple replant disease soil; -ARD: gamma-irradiated ARD soil; Control: soil with no apple history). We investigated the microbial community composition by 16S rRNA gene amplicon sequencing (bacteria and archaea) along the soil-root continuum (bulk soil, rhizosphere, rhizoplane). Significant differences in microbial community composition and structure were found between +ARD and -ARD or +ARD and Control along the soil-root continuum, even for plants exposed simultaneously to two different substrates (-ARD/+ARD, Control/+ARD). The substrates in the respective split-root compartment defined the assembly of root-associated microbial communities, being hardly influenced by the type of substrate in the respective neighbor compartment. Root-associated representatives from Actinobacteria were the most dynamic taxa in response to the treatments, suggesting a pivotal role in ARD. Altogether, we evidenced an altered state of the microbial community in the +ARD soil, displaying altered alpha- and beta-diversity, which in turn will also impact the normal development of apple rhizosphere and rhizoplane microbiota (dysbiosis), concurring with symptom appearance.

Pore structure is a key determinant of soil functioning, and both root growth and activity of soil fauna are modified by and interact with pore structure in multiple ways. Cover cropping is a rapidly growing popular strategy for improving agricultural sustainability, including improvements in pore structure. However, since cover crop species encompass a variety of contrasting root architectures, they can have disparate effects on formation of soil pores and their characteristics, thus on the pore structure formation. Moreover, utilization of the existing pore systems and its modification by new root growth, in conjunction with soil fauna activity, can also vary by cover crop species, affecting the dynamics of biopores (creation and demolition). The objectives of this study were (i) to quantify the influence of 5 cover crop species on formation and size distribution of soil macropores (>36 μm Ø); (ii) to explore the changes in the originally developed pore architecture after an additional season of cover crop growth; and (iii) to assess the relative contributions of plant roots and soil fauna to fate and modifications of biopores. Intact soil cores were taken from 5 to 10 cm depth after one season of cover crop growth, followed by X-ray computed micro-tomography (CT) characterization, and then, the cores were reburied for a second root growing period of cover crops to explore subsequent changes in pore characteristics with the second CT scanning.Our data suggest that interactions of soil fauna and roots with pore structure changed over time. While in the first season, large biopores were created at the expense of small pores, in the second year these biopores were reused or destroyed by the creation of new ones through earthworm activities and large root growth. In addition, the creation of large biopores (>0.5 mm) increased total macroporosity. During the second root growing period, these large sized macropores, however, are reduced in size again through the action of soil fauna smaller than earthworms, suggesting a highly dynamic equilibrium. Different effects of cover crops on pore structure mainly arise from their differences in root volume, mean diameter as well as their reuse of existing macropores.