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Earthworm Burrowing Activity and Its Effects on Soil Hydraulic Properties

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Earthworm Burrowing Activity and Its Effects on Soil Hydraulic Properties sustainability Article Earthworm Burrowing Activity and Its Effects on Soil Hydraulic Properties under Different Soil Moisture Conditions from the Loess Plateau, China 1,2 1,2,3, , 1, , Shuhai Wen , Ming’an Shao * y and Jiao Wang * y 1 Key Laboratory of Ecosystem Network Observation and Modelling, Institute of Geographic Sciences and Natural Resources Research, Chinese Academy of Sciences, Beijing 100101, China; [email protected] 2 College of Resources and Environment, University of Chinese Academy of Sciences, Beijing 100190, China 3 State Key Laboratory of Soil Erosion and Dryland Farming on the Loess Plateau, Northwest A&F University, Yangling 712100, China * Correspondence: [email protected] (M.S.); [email protected] (J.W.) Present address: 11A, Datun Road, Chaoyang District, Beijing 100101, China. y Received: 15 September 2020; Accepted: 4 November 2020; Published: 9 November 2020 Abstract: Earthworm activity has become more important in the Loess Plateau, where hydrological processes are crucial for ecosystem sustainability. In this study, we conducted a laboratory microcosm experiment to determine the various burrowing activities of Eisenia fetida and their impact on the soil hydraulic properties in response to different levels of soil moisture (50%, 70%, 90% of field capacity) in two common soil types (loessial and Lou soil) obtained from the Loess Plateau. Burrowing activity of E. fetida increased with higher soil moisture and was greater in loessial than in Lou soil. Most burrowing activities occurred within the top 5 cm and decreased with increasing soil depth. Macropores and burrow branching, which are highly related to the earthworm burrowing, were more prevalent in wetter soil. Earthworms significantly altered the formation of large soil aggregates (AGL, diameter >2 mm) under different soil moistures and depths. Distinct earthworm burrowing activities, controlled by soil moisture, altered soil hydraulic properties. However, soil saturated hydraulic conductivity (Ks) showed little differences between different treatments due to the horizontal and high–branched burrows of E. fetida, although higher burrowing activities were found in wetter soil. Soil field capacity was highest in drier soil due to the less macropores and burrowing activities. Keywords: earthworm burrows; soil moisture; soil aggregates; hydraulic properties; Loess Plateau 1. Introduction Earthworms play an important role in the regulation of soil physicochemical processes and ecosystem services, such as the maintenance of soil structure, water regulation, and nutrient inputs for plants [1,2]. Their burrowing activities increase the proportion of macropores in the soil, which have important hydrological functions: (1) preferential flow through macropores could improve water movement to the deep soil, enhance water infiltration and drainage, reduce surface runoff, and subsequently, decrease soil erosion [3,4]; (2) increased soil aeration in the macropores enhances water loss through evaporation, decreasing soil moisture content [5]. However, the variability in earthworm burrowing activities is strongly related to soil properties (e.g., soil type, soil texture, organic matter) and soil temperature and is particularly sensitive to soil moisture [6]. Several studies have shown that earthworms were often inactive, entered diapause, and lost weight when soil moisture was low [7,8]. Limited burrowing activities were observed in some laboratory experiments when the soil water potential or water content decreased [9–11]. A wet habitat is well–known to be essential for earthworm activity; however, Perreault and Whalen [12] found greater earthworm burrowing Sustainability 2020, 12, 9303; doi:10.3390/su12219303 www.mdpi.com/journal/sustainability Sustainability 2020, 12, 9303 2 of 13 activity in drier soil than wetter soil, as earthworms explore a larger volume of soil for their survival when the soil is dry. Earthworm activity is based on complex interactions between soil conditions, climate, and earthworm species and has been well documented in Europe and North America for decades [13,14]. Thus, more basic studies evaluating the effect of soil conditions (e.g., moisture) on burrowing activity are critically needed in soils of Asia, which has received little attention. Earthworms form a mass of soil aggregates through casting and burrowing, which could account for 50% of the topsoil aggregates in temperate regions [6,15]. Earthworms redistribute fine particles within the soil by aggregation, wherein the advanced large pores between aggregates and inner pore structure of casts (bio–aggregates) can improve infiltration, water retention, and discharge [16–19]. Some studies have shown that aggregation due to casting activities is related to soil moisture [6,20]. A model developed based on earthworm biomass, soil moisture, and temperature estimated that the soil aggregate formation rate by earthworms increased with higher soil water potential [21]. Hindell et al. [22] found that earthworm casting activity was greatest at a high soil water potential of 0 to 2 kPa in a gradient of low to high (from 25 kPa to 2 kPa) water potential. − − − The impact of earthworms on soil hydraulic properties is well–studied by describing the burrowing activity and burrow characteristics [23–25]. Higher permeation rates and faster water discharge were observed resulting from the high burrowing activities of earthworms. In addition, soil Ks increased as more surficial, connected, and continuous burrows are created by earthworms [23,24,26]. However, horizontally created earthworm burrows with many branches have been found to limit or reduce the water conductivity [27,28], indicating a variety of water–based processes determined by earthworm burrowing activities. Water retention is therefore closely related to the earthworm activities. Indeed, water holding capacity can decrease with increasingly efficient water drainage or faster evaporation through earthworm burrows [2]; conversely, it can increase following the increased content of water–stable aggregates also formed by earthworms [29]. Since soil moisture is one of the most important factors influencing the earthworm activities, the impact of earthworms on soil hydraulic properties may vary depending on soil moisture. Therefore, the relationship between the various burrowing activities and their effects on water transfer, water–retention properties under different soil moisture levels merit investigation. The Loess Plateau is widely known for having severe soil erosion and hydrological problems; hence, it is urgent to evaluate and improve the efficiency of water infiltration and conservation in this area [30]. Soil macrofauna has already been shown to play an important role on soil hydrological processes in the region. A series of studies by Li et al. [31,32] reported the importance of crickets and ants on regulating soil structure and water movement on the Loess Plateau. The burrows of mole crickets effectively intercepted rainfall, improved water infiltration, and reduced runoff; the preferential flow into the burrows and nests also effectively increased soil moisture content. There is a surprising lack of research on earthworms on the Loess Plateau, even though earthworms are one of the major soil engineers and are used frequently in ecological farming in recent years. E. fetida is a common species used in ecological farming for its prominent capacity for organic matter decomposition and vermicomposting [33]. Therefore, most studies on E. fetida have been carried out to investigate biodegradation and nutrient recycling. Despite the species’ intense burrowing activity within the topsoil layer, impacts on soil’s physical and hydraulic properties are less considered and therefore necessary to evaluate the role of earthworms on ecological sustainability on the Loess Plateau. Here, we conducted a microcosm experiment to quantitatively characterize the burrows of E. fetida under three different levels of soil moisture in two common soil types from the Loess Plateau. In addition, soil aggregation and hydraulic properties were determined to evaluate the effects of the burrowing activity of E. fetida on soil physical–hydrological functions. Sustainability 2020, 12, 9303 3 of 13 2. Materials and Methods 2.1. Experimental Design The study was conducted based on microcosm experiment using a completely randomized factorial design with two soil types: Lou soil and loessial soil, and under three soil moisture conditions: 50%, 70%, and 90% of field capacity. A native and dominant epigeic earthworm species E. fetida was used for the study. Lou soil and loessial soil are two common soil types occurring on the central and southern part of the Loess Plateau, respectively. Lou soil is clay loam, described as a Cumulic Anthrosol and loessial soil is silty clay loam (FAO/UNESCO); physical and chemical properties of the soil types are shown in Table S1. Lou soil was obtained from the upper 20 cm of a wheat–fallow field in Yangling (108◦040 E, 34◦170 N), while loessial soil was obtained from the upper 20 cm of an apple orchard in Changwu (107◦410 E, 35◦140 N). The soil was air–dried to a soil moisture content of less than 5% and sieved at 2 mm. The microcosms used in this study were plexiglass columns with diameter of 20.4 cm and height 3 3 of 25 cm. Soil was packed to a height of 20 cm at a dry bulk density of 1.19 g cm− and 1.16 g cm− for Lou and loessial soil, respectively; the density of each soil type was established by a field test conducted at each study site. In this process, the soil in each column was compacted stepwise in three layers to minimize the variation in the bulk density within the soil column. Mesh fabric was placed at the top and bottom of the column to prevent earthworms escaping and soil loss. A plastic tray was placed at the bottom of the column for water supplementation. E. fetida, a widely distributed species in both soil types, was collected from the study sites. The average weight of an individual was about 0.5 g. Six adult earthworms were introduced into each microcosm, and 5 g of litter and 100 g of cow manure were added to the soil surface as the food supply. After the earthworms were added, soil moisture was supplied at three different levels.
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