Aim We investigated how substrate hydraulic properties respond to the presence of arbuscular mycorrhizal fungi (AMF) in root-containing and root-free substrate zones in a Medicago truncatula-Rhizophagus irregularis model system. Methods Before planting, two compartments constructed from standard soil sampling cores (250 cm3) were implanted into non-mycorrhizal and mycorrhizal pots containing a sand-zeolite-soil mix. One compartment allowed root penetration (1 mm mesh cover) and the other only hyphal ingrowth (42 μm mesh cover). After eight weeks of growth under maintenance of moist conditions, the cores were subjected to water retention measurements. Additionally, we measured water retention of bare substrates before and after drying events to check for successful maintenance of moist conditions in pots. Results Drying of bare substrates decreased water retention, but planting at least sustained it. The parameters of water retention models responded linearly to root morphological traits across mycorrhizal and non-mycorrhizal substrates. Hyphae-only colonization comparatively affected the course of water retention in ways that suggest increased pore space heterogeneity while maintaining water storage capacity of substrates. Conclusions Hence, water contents corresponded to different substrate matric potentials in non-mycorrhizal and mycorrhizal pots. We conclude that changes to water retention in AMF colonized substrates can contribute to a widely observed phenomenon, i.e. that mycorrhizal plants differ in their moisture stress response from non-mycorrhizal plants.

Arbuscular mycorrhizal fungi (AMF) alter plant water relations and contribute to soil structure. Although soil hydraulic properties depend on soil structure and may limit plant water uptake, little is known about how AMF influence soil water retention (the relation between the soil water content and soil water potential) and hydraulic conductivity in different soils. Instead, these soil hydraulic properties often are considered to be independent of AMF presence in experiments. We asked if this assumption holds true for both sand and loam. We grew maize plants either inoculated with Rhizophagus irregularis or with autoclaved inoculum in pots filled with quartz sand or loam soil until extraradical spread of the fungus throughout the pots was achieved. Each pot contained a hyphal compartment made of a soil sampling core (250 cm3) covered with a 20-µm nylon mesh to encourage fungus ingrowth but to exclude root ingrowth. We measured soil water retention and unsaturated hydraulic conductivity in these undisturbed root-free soil volumes. We observed that in loam harboring the mycorrhizal fungus, the soil water retention decreased, while in sand, it increased without detectable changes in the soil bulk density. The effects of the fungus on the soil water potential were strongest at low soil water contents in both soils. As a consequence of the altered water potentials in soils with the mycorrhizal fungus, soil hydraulic conductivity increased in loam but decreased in sand after fungus ingrowth. We conclude that in our study, the mycorrhizal fungus acted as a soil conditioner even distant from roots, which encouraged drainage in loams prone to sogginess but enhanced water storage in sands prone to quick desiccation. We recommend considering soil hydraulic properties as being dynamic in future studies on water relations of mycorrhizal plants.

<p>Hydraulic properties of mycorrhizal soils have rarely been reported and difficulties in directly assigning potential effects to hyphae of arbuscular mycorrhizal fungi (AMF) arise from other consequences of AMF being present, i.e. their influence on growth and water consumption rates of their host plants that both also influence soil hydraulic properties.</p><p>We assumed that the typical nylon meshes used for root-exclusion experiments in mycorrhizal research can provide a dynamic hydraulic barrier. It is expected that the uniform pore size of the rigid meshes causes a sudden hydraulic decoupling of the enmeshed inner volume from the surrounding soil as soon as the mesh pores become air-filled. Growing plants below the soil moisture threshold for hydraulic decoupling would minimize plant-size effects on root-exclusion compartments and allow for a more direct assignment of hyphal presence to modulations in soil hydraulic properties.</p><p>We carried out water retention and hydraulic conductivity measurements with two tensiometers introduced in two different heights in a cylindrical compartment (250 cm³) containing a loamy sand, either with or without the introduction of a 20 µm nylon mesh equidistantly between the tensiometers. Introduction of a mesh reduced hydraulic conductivity across the soil volumes by two orders of magnitude from 471 to 6 µm d<sup>-1</sup> at 20% volumetric water content.</p><p>We grew maize plants inoculated or not with Rhizophagus irregularis in the same soil in pots that contained root-exclusion compartments while maintaining 20% volumetric water content. When hyphae were present in the compartments, water potential and unsaturated hydraulic conductivity increased for a given water content compared to compartments free of hyphae. These differences increased with progressive soil drying.</p><p>We conclude that water extractability from soils distant to roots can be facilitated under dry conditions when AMF hyphae are present.</p><p> </p>