Applied Soil Ecology 134 (2019) 31–37

Contents lists available at ScienceDirect

Applied Soil Ecology

journal homepage: www.elsevier.com/locate/apsoil

Soil compaction effects on litter decomposition in an arable field: T

Implications for management of crop residues and headlands
Lea Carlessoa, , Andrew Beadleb, Samantha M. Cookc, Jess Evansc, Graham Hartwelld, Karl Ritze,
⁎

Debbie Sparkese, Lianhai Wua, Phil J. Murraya

a
Rothamsted Research, North Wyke, Okehampton EX20 2SB, UK
b
BASF SE, APD/S, 67117 Limburgerhof, Germany
c
Rothamsted Research, Harpenden, Hertfordshire AL5 2JQ, UK
d
BASF Environmental Stewardship & Crop Protection, Cheadle SK8 6QG, UK
e
University of Nottingham, Sutton Bonington, Leicestershire LE12 5RD, UK

ARTICLE INFO ABSTRACT

Keywords: Soil compaction is a major threat to agricultural soils. Heavy machinery is responsible for damaging soil che-
Decomposition mical, physical and biological properties. Among these, organic matter decomposition, which is predominantly
Compaction mediated by the soil biota, is a necessary process since it underpins nutrient cycling and the provision of plant
Field margins nutrients. Understanding factors which impact the functionality of the biota is therefore necessary to improve
Environmental stewardship scheme
agricultural practices. To better understand the effects of compaction on the soil system, we determined the
Soil quality
effects of soil bulk density and soil penetration resistance on the decomposition rates of litter in three distinct
field zones: a grass margin, sown at the edge of the field adjacent to the crop, tramlines in the crop:margin
interface, and crop. Three litters of different quality (ryegrass, straw residues and mixed litter) were buried for 1,
2, 4 and 6 months in litter bags comprising two different mesh sizes (0.02 and 2 mm). Bulk density and soil
penetration resistance were greater in the compacted tramline than in the margin or the crop. The greatest
amount of litter remaining in the bags after 6 months was found in the tramline, and the least in the grass
margin. Differences between treatments increased with burial time. No significant differences in mass loss be-
tween the two mesh sizes was detected before the fourth month, implying that microbial activities were the main
processes involved in the early stages of decomposition. Decomposition in the tramline was clearly affected by
the degradation of soil structure due to heavy compaction. This study shows that soil conditions at the edges of
arable fields affect major soil processes such as decomposition. It also reveals the potential to mitigate soil
degradation by managing the headland, the crop residues and the machinery traffic in the field.

1. Introduction 2014). This included compensation for the setting up of grass margins
around arable fields with the primary aim of encouraging aboveground
Land-use is a primary determinant in driving soil processes (Holland biodiversity (Department for Environment, Food and Rural Affairs,
et al., 2014; Postma-Blaauw et al., 2010; Sousa et al., 2004). It has been 2014; Meek et al., 2002). Evidence suggests such margins can provide
shown that vegetation cover modifies soil biodiversity (Crotty et al., important ecosystem services including pollination and pest manage-
2015, 2014) and that the more intense the land use (intensive crop ment (e.g. Lu et al., 2014). However, the implications for the below-
production in comparison to extensive grassland), the fewer the number ground biodiversity and the functions they support have been con-
of functional groups prevail (Tsiafouli et al., 2015). In 1994, the United sidered less, even though it has been shown that the soil biota can be
Kingdom government published a Biodiversity Action Plan, establishing adversely affected by field management (Sechi et al., 2017). Field
arable field margins at the edge of fields as priority habitat (Maddock, margins affect nutrient transformation and run-off (Marshall and
2008) and supported by a new environmental stewardship scheme for Moonen, 2002), and the soil fauna plays a pivotal role in many of the
farmers to increase and support biodiversity in the agricultural land- soil processes that, in turn, deliver ecosystem services (Bardgett and van
scape in 2014 (Department for Environment, Food and Rural Affairs, der Putten, 2014; Wall et al., 2015). Among these services,

⁎
Corresponding author at: Sustainable Agriculture Sciences, Rothamsted Research, North Wyke, Okehampton EX20 2SB, UK.
E-mail address: lea.carlesso@rothamsted.ac.uk (L. Carlesso).

https://doi.org/10.1016/j.apsoil.2018.10.004
Received 9 July 2018; Received in revised form 4 October 2018; Accepted 6 October 2018
Available online 25 October 2018
0929-1393/ © 2018 The Authors. Published by Elsevier B.V. This is an open access article under the CC BY license
(http://creativecommons.org/licenses/BY/4.0/).
L. Carlesso et al. Applied Soil Ecology 134 (2019) 31–37

decomposition, a biologically-driven process, underpins nutrient cy- agricultural systems and to mitigate the impacts of compaction.
cling and primary production (Coleman et al., 2004; Hättenschwiler In this study, we determined organic matter decomposition rates of
et al., 2005). The interaction between different classes of organisms plant material (wheat straw and ryegrass residues) in contrasting zones
(microbiome and macrobiome) is necessary to ensure the decomposi- of an arable field that had been subjected to different pressures. We
tion of primary organic matter (Bradford et al., 2002). Although the aimed to identify effects of machinery wheeling and agricultural man-
role of the microbiome (bacteria, archaea and fungi) is reasonably well agement on decomposition and understand how the response changes
understood, Setälä et al. (1996) demonstrated the benefits of a more with respect to litter type and soil faunal exclusion. We hypothesised:
complex community for improved nutrient cycling. It has also been (i) decomposition rate would be lowest in more compacted soils; (ii)
shown that macrofauna modify the processes of decomposition by their ryegrass litter, because of its lower C:N ratio, would decompose faster
action on the microbiota (Hättenschwiler et al., 2005; Joly et al., 2015). than straw residues, the decomposition rate of an equal mixture of both
Due to it’s impact on organic matter dynamics (Wolters, 2000), the litters would fall within the interval between the two; (iii) exclusion of
presence of the macrobiome (meso- and macrofauna) is required to the soil meso- and macrofauna would reduce the decomposition rate.
achieve the decomposition of plant litter and therefore should be re-
garded as a potential tool for crop management and nutrient cycling in
agricultural contexts. In an such contexts, most of the litter decom- 2. Materials and methods
position is affected by agronomic practices, such as management of
crop residues, fertilisation or soil compaction from agricultural ma- 2.1. Site and soil characteristics
chinery. The amount and quality of organic matter returned to the
system (Fierer et al., 2005; Gergócs and Hufnagel, 2016; Milcu and The experiment was carried out between October 2016 and April
Manning, 2011) together with the presence of faunal and microbial 2017 at The Grange Farm, Northamptonshire, United Kingdom (52° 18′
communities (Murray et al., 2009) are primary factors regulating de- 2.73″ N; 0° 45′ 52.83″ W) in an arable field planted with oilseed rape
composition rates. Thiele-Bruhn et al. (2012) noted the capability of (Brassica napus L.) which had previously been in winter wheat (Triticum
agricultural practice to control the quality of primary organic matter aestivum L.). The field had been managed using minimum tillage tech-
entering soil systems and therefore its capacity to modify the soil niques (i.e. no deep ploughing) for at least 15 years. Mineral fertilisa-
community and its activity. To understand the effects of litter quality, tion and chemical inputs were applied to the crop following the UK
Johnson et al. (2007) tested the decomposition of five crops of varying standard scheme management for farmers (Agriculture and
chemical composition and three different organs of each plant, and Horticulture Development Board, 2017). The crop was planted in a field
showed that crop and plant parts affected decomposition rates and C- bordered by a 6 m-wide 10-year-old grass margin that had been set up
pools at the soil surface. This implies some potential for agricultural soil to promote biodiversity in the agricultural landscape (Department for
management via crop residues. Environment, Food and Rural Affairs, 2014; Maddock, 2008). The soil
The architecture of the habitat and the associated propensity for was classified as Hanslope series, a typical calcareous pelosol from a
belowground oxygen supply (modulated by the soil pore networks) are clayey chalky drift series (calcaric stagnic cambisol soil) with poor
two more factors affecting decomposition rates. The deterioration of drainage capacity and high sensitivity to compaction (Cranfield
soil structure (principally via a reduction in porosity and connectivity University, 2017).
of pores) caused by external factors has been shown to affect microbial During the period of study climate conditions were characterised by
mineralisation (Beylich et al., 2010; De Neve and Hofman, 2000), as two dry periods; one in October at the beginning of the experiment and
well as habitat and food resources that support the soil fauna (Beylich one in December. Temperatures were normal for the region (Fig. S1).
et al., 2010; Althoff et al., 2009; Larsen et al., 2004). In agricultural The experimental area consisted of 18 plots (6 × 6 m) distributed
landscapes, soil structure is exposed to deterioration by heavy ma- among six blocks along the south side of the field. Each block comprised
chinery traffic and many arable soils are sensitive to increased com- three plots, one in each of three field ‘zones’: one in the grass margin,
paction, causing a decline in crop yield (Hamza and Anderson, 2005). one in the tramlines between the margin and the crop, which were
Within the scope of environmental schemes, and to prevent damage to visibly compacted, and one in the actual crop. The field zones (margin,
improved biodiversity habitats such as field margins, the policy re- margin-crop interface and crop) were spatially constrained and so
quires that farmers do not manoeuvre on the field margins, obliging randomisation of plots within blocks was not possible. Total soil carbon
them to turn at the edges of the crop and thus creating a compacted (C) and total soil nitrogen (N) concentrations were measured using an
area between the main crop and the margin. A better understanding of elemental analyser (N1500, Carlo Erba, Milan, Italy). C:N ratio was
the effects of compaction on organic matter decomposition and biolo- determined as average values calculated from cores taken at each of the
gical activity in soils is a necessary step to improve soil management in 18 plots.

Table 1
Average values (n = 6) and standard error ( ± SE in italic) of various soil properties measured in three zones of an oilseed rape field (October 2016). Superscript
letters a and b show significant difference of means between the field zones (Least square difference test, Bonferroni adjustment).
Field Water content (% Bulk density (g.cm−3) Total C (% Volume of Total N (% Volume of C:N ratio
zones Volume of Soil−1) Soil−1) Soil−1)

Grass margin
Mean 17.4a 0.89a 4.08a 0.40a 10.20a
± SE 1.02 0.03 0.39 0.02 0.39

Tramline
Mean 11.7b 1.25b 2.32b 0.27b 8.61b
± SE 0.62 0.03 0.14 0.01 0.46

Crop
Mean 14.6a 1.02a 2.36b 0.25b 9.58b
± SE 0.25 0.06 0.19 0.02 0.42

32
L. Carlesso et al. Applied Soil Ecology 134 (2019) 31–37

2.2. Soil compaction assessment dried and weighed as described above. The proportion of litter re-
maining following each burial duration was then calculated.
Soil bulk density (Laryea et al. (1997) was determined from cores
(8 cm diameter × 10 cm depth) taken at random from each of the 18 2.4. Statistical analyses
plots at the beginning of the experiment. This sampling method was
considered appropriate for our requirements as it has been shown to not Impacts of field zone on bulk density was estimated by a 1-way
significantly affect bulk density measurement (Özgöz et al., 2006; Page- analysis of variance (ANOVA). An analysis of covariance (ANCOVA)
Dumroese et al., 1999). Samples were dried at 105 °C for 24 h, then was used to test the effect of field zone (grass margin, tramline and
plant residues and stones were removed by 2 mm sieving. Water con- crop) on soil penetration resistance, controlling for the effects of depth,
tent was calculated from the proportion of dry soil to wet soil (Table 1). which co-vary with the field zone effect. We tested the similarity of soil
Soil penetration resistance was recorded on April 1st 2017 with a pe- penetration resistance in the “grass margin” and the “crop” zone by
netrometer (Solutions for Research Ltd, Silsoe, Bedfordshire, UK) fitted using a nested treatment structure in the model; the ANCOVA therefore
with a 9.45 mm diameter (base area 7 × 10−5 m2), 30-degree cone. At the effect of the uncompacted zones (grass margin and crop) versus the
every sampling point, the soil penetration resistance was measured at compacted tramline as well as comparing the effect of grass margin
14 depth points each 3.7 cm apart (from 3.7 cm to 51.8 cm depth). versus crop.
Penetrometer resistance was calculated by dividing force at each depth The five samples taken from the initial litter material (T0) are
by the cone base area. Ten replicate measurements were randomly pseudoreplicates in terms of testing for treatment effects and as such
taken at each plot. Data were calibrated by Solutions for Research Ltd, were not analysed statistically. However, means and standard errors
Silsoe, Bedfordshire, UK; and soil strength (Pa) was determined from have been presented as a basis for comparison between litter types and
the force measured by the penetrometer (kg × 9.81), divided by the for comparing later time points to baseline. A two-way split plot ana-
base area (m2), so that a 10 kg force reading represented 1399 KPa. lysis of variance (ANOVA) was used to test for the differences of C:N
ratio between two treatment factors (mesh size and litter type) and
2.3. Organic matter decomposition experiment their interaction. We used a post-hoc comparison Tukey test at 95%
confidence level to see where differences between factors occurred.
Litter bags (6 cm length × 5 cm height) were made using two mesh A four-way split plot ANOVA was used to determine effect of the
sizes; one set with a plastic mesh size of 2 mm allowed full access by the treatment factors (mesh size, litter type, field zone and time period in
soil biota, and one set with 0.02 mm nylon mesh which excluded most the ground) and their interactions on the quantity of litter remaining at
of the fauna and allowed microbial access only. the end of the experiment. Because of the destructive sampling of the
Three types of litter of different quality (C:N ratio) were prepared: a litter bags, time was not considered as a repeated measurement. We
low C:N ratio perennial ryegrass (Lolium perenne L.), a high C:N ratio used Fisher’s Least significant difference method (LSD) to assess whe-
wheat straw (T. aestivum) and a 50:50 mixture of both types of litter of ther pairwise combinations were different from one another, with
intermediate C:N ratio. Ryegrass and wheat straw were oven dried to Bonferroni adjustment. Similarly, the effect of mesh size, litter type and
constant weight at 105 °C. Straw and ryegrass were chopped into ap- field zone were analysed using a 3-way ANOVA for month 1, 2, 4 and 6
proximately 1.5–2 cm length pieces, then, 1.0 g of the litter was added separately. All statistical analyses were done using R software 3.1.2
to each of the litter bags (0.5 g of both chopped litter types was added (http://www.r-project.org/).
for the mixed litter treatment). Litter bags were carefully stored in in-
dividual boxes to prevent physical damage during transportation; litter
loss (collected in the bottom of boxes after transportation) was con- 3. Results
sidered insignificant in terms of affecting future measurement. Average
values of total carbon and total nitrogen of wheat straw and ryegrass 3.1. Soil compaction
were measured from 5 subsamples of each of the initial materials (i.e.
T0), and after 6 months (T6) from material remaining in both the small Bulk density was significantly greater in the compacted area of the
and large mesh bags, for each of the three litter types (ryegrass, straw tramline compared to either the grass margin or the crop (1-way
and a 50:50 mixture of both) using an elemental analyser (N1500, Carlo
Soil penetration resistance (KPa)
Erba, Milan, Italy). The initial C:N ratio of the 50:50 mixture was taken 0 1000 2000 3000 4000 5000 6000
as the arithmetic mean of the constituent ratios. 0
In total, 432 litter bags were prepared, half with the small mesh size
and half with the large mesh. We allocated 24 litter bags to each of the 10
18 plots (four bags of each mesh size containing straw, rye or the 50:50
mixture) and buried these on 1 October 2016. A sub-set of 108 bags
(one bag of each treatment combination from every plot) were removed 20
Depth (Cm)

on 1 November 2016, 1 December 2016, 1 February 2017 and 1 April

2017, representing 1, 2, 4 and 6 months’ burial duration. The latter time 30
is consistent with the cropping cycle, when the cultivated part of the
field is then physically disrupted. Litter bags were buried in the top soil
at 5 cm depth in each plot and the position of bags of each treatment 40

combination was completely randomised within the plot. To minimise

disturbance and to preserve the context of the inherent soil structure as 50
far as possible, a vertical slit was made with a spade, just sufficient to
locate the bag, and then closed up by firming the soil back into position.
Tramline Crop Grass margin
A string and a knot code system were used to identify each treatment.
One bag was missing on the first and third collection dates, and 5 bags Fig. 1. Profiles of soil penetration resistance (KPa) within three field zones
were missing on the last date. (grass margin ■, tramline wheeling in the crop-margin interface ● and, crop
After removing the litter bags from the ground, the litter was re- ▲) at 14 depth points (3.7–51.8 cm depth) within a field containing oilseed
moved from the bags; soil particles were gently washed away from the rape, 2017 cropping season. Points show means (n = 60); bars denote standard
litter using a 15 μm sieve to retain plant materials. The litter was then error. In some instances, these fall within the confines of the symbols.

33
L. Carlesso et al. Applied Soil Ecology 134 (2019) 31–37

100 100 100

90 90 90
80 80 80
Litter remaining (%)

70 70 70
60 60 60
50 50 50
40 40 40
30 30 30
20 20 20
10 10 10
0 0 0
0 1 2 3 4 5 6 0 1 2 3 4 5 6 0 1 2 3 4 5 6

a Time (month) Time (month) Time (month)

b c
Large
Small

Ryegrass 50 :50 Mix Straw

Fig. 2. Percentage of different litter types (perennial ryegrass (Lolium perenne) black ●, wheat straw (Triticum aestivum) light grey ■, and a 50:50% mixture of both
litters grey▲) remaining after 1, 2 ,4 and 6 months buried in three different zones of a field containing oilseed rape, in litter bags with small (0.02 mm) and large
(2 mm) mesh sizes; Zones: grass margin (2a), compacted tramline (2b) and crop (2c)). Month 0 corresponds to the start of the experiment (1st October 2016) and
Month 6 the end of the experiment (1st of April 2017). Points show means (n = 18); bars denote standard error. (For interpretation of the references to color in this
figure legend, the reader is referred to the web version of this article.)

ANOVA F(2,13) = 18.6, P < 0.001; Table 1). 90

The soil penetration resistance increased significantly with depth in 80

all of the three field zones (ANCOVA F(1,246) = 1003, P < 0.001). A
peak was observed at 7.4 cm in the tramline, whereas the slope of the 70

resistance in the crop increased below the historical ploughed layer at 60

23 cm depth (Fig. 1). There was no difference in soil penetration re-
sistance between the crop and the field margin zones (ANCOVA
C:N ratio

50

F(1,246) = 0.23, P = 0.63) and overall, the soil penetration resistance 40

was significantly greater in the compacted zone (tramline in the crop-
margin interface) than in the uncompacted zones (crop and field margin 30

zones combined) (ANCOVA F(1,246) = 129, P < 0.001). 20

10
3.2. Litter decomposition
0
Large Small Large Small Large Small
3.2.1. Comparison of the two mesh sizes
Gras s Mix Straw
In the first two months of the experiment, regardless of the field
Litter type and mesh size treatment
zone or the litter type, there was no significant difference in litter re-
maining between the large and small mesh size bags (3-way ANOVA for Fig. 3. Carbon:nitrogen ratios of the three plant residues used in litter bags of
Month 1 and Month 2, F(1,74) = 0.63, P = 0.431 and F(1,75) = 0.67, large (2 mm) and small mesh size (0.02 mm) at the outset of the study (T0), and
P = 0.415, respectively). However, from Month 4, there was generally main effects of litter type and mesh size after 6 months burial (T6). The grey bar
more litter remaining undecomposed in the small mesh than in the charts display C:N values at T6 (error bar showed standard error), while the
large mesh size bags (3-way ANOVA F1,74 = 69.3, P < 0.001; Fig. 2). black dots and the broken lines denote C:N ratios at T0. For the mixture, this is
calculated as arithmetic mean of ryegrass and straw values based upon a
This effect was persistent at Month 6 (3-way ANOVA F(1,70) = 92.7,
50:50% mix of these constituents. (For interpretation of the references to color
P < 0.001). Overall the combined effect of mesh size on litter re-
in this figure legend, the reader is referred to the web version of this article.)
maining over time was significant (4-way ANOVA F(3,350) = 34.84,
P < 0.001). The effect of the field zone combined with the mesh size
was also significant (4-way ANOVA F(2,350) = 3.65, P = 0.027), with 47.1 ± 8.1% of the ryegrass was left. Mixed litter had an intermediate
relatively less litter lost in the large compared to the small mesh bags decomposition rate, with 60.4 ± 8.0% of material remaining. There
when these were buried in the tramline or the crop rather than in the was a significant interaction of the combined effects of litter type and
grass margin zone (Fig. 2). mesh size (4-way ANOVA F(2,350) = 22.4, P < 0.001), with less rye-
grass litter remaining in the large litter bags than straw residues or
mixed litter in small and large mesh size bags (Fig. 2). Even though the
3.2.2. Effects of crop litter quality difference in litter remaining between the two mesh sizes at Months 1
Litter type significantly affected the proportion of plant material and 2 was not significant, remaining ryegrass litter in the large mesh
remaining in the bags at the end of the experiment (4-way ANOVA size litter bags was marginally smaller than in other treatments (3-way
F(2,350) = 386, P < 0.001); A mean across treatments ± standard ANOVA F(2,74) = 3.08, P = 0.052 and F(1,75) = 2.94, P = 0.059 for
error (SE) of 72.6 ± 7.3% of the straw remained after 6 months, while

34
L. Carlesso et al. Applied Soil Ecology 134 (2019) 31–37

Month 1 and Month 2, respectively; Fig. 2). would be exceed and consequently the limited supply of oxygen would
The C:N ratio of the litter declined over time only in the case of restrict decomposition processes (Beylich et al., 2010; Horn et al., 1995;
straw and the straw:ryegrass mixture (Fig. 3). After 6 months burial Whalley et al., 1995). The shallow angle of the slope observed in the
duration, there was no significant interaction between litter type and soil penetration resistance measurements from the cropped zone cor-
mesh size, but a highly significant main effect due to litter type (2-way responds to the historical ploughed layer at 23 cm. Above this layer, soil
ANOVA F(2,80) = 156, P < 0.001), where the proportion of difference penetration resistance in the grass margin and the crop zone behaved
between T0 and T6 was around three-fold greater in the case of straw differently but reached similar intensities below this interface. Even
compared to ryegrass (Fig. 3). In the case of the mixture, the C:N ratio though the field had been farmed using minimum tillage cultivation
was about half that of the straw, but significantly greater than that for techniques for the past 15 years, this could reflect the long-term effect
ryegrass alone (Fig. 3). There was also a significant effect of mesh size of previous ploughing practices on soil structure. The potential impact
upon C:N ratio after 6 months where the ratio was 10% smaller in small of this on soil processes (e.g. Peigné et al., 2013) warrant further in-
compared to large mesh bags (2-way ANOVA F(1,75) = 5.30, P = 0.02; vestigation. Our results showed that decomposition occurs more slowly
Fig. 3). in the compacted soil of the tramlines at the crop-margin interface re-
gardless of the litter type or the mesh size of the bags used in the ex-
3.2.3. Effect of the field zone on litter losses periment.
The location of the litter bags in the field (zone) significantly af- The two different mesh sizes of litter bags used in the decomposition
fected the decomposition rate of all litter types within bags of the two experiment enabled conclusions to be drawn about the effects of mi-
different mesh sizes (4-way ANOVA F(2,10) = 34.0, P < 0.001). With a crobial communities and larger soil fauna on decomposition since the
mean of 64.8 ± SE 7.8%, remaining litter was greater in bags placed in large mesh size allowed access of all soil fauna and the small mesh size
the tramline and similar decomposition rates were observed in bags excluded this fraction and would therefore be predominantly microbial
buried in the grass margin and in the crop (on average 55.0% and (Bokhorst and Wardle, 2013). Before Month 4, there was no difference
60.2% of litter remaining, respectively). in mass loss between litter bags of the two mesh sizes, implying that
initial decomposition (Month 1 and Month 2) was primarily carried out
4. Discussion by microbes or that effect of the soil fauna was negligible. In this study,
the addition of an exogenous source of organic matter might have sti-
We hypothesised that deteriorated soil conditions, caused by traf- mulated primary microbial colonisation, resulting in mineralisation of
ficking, would reduce plant litter decomposition at the interface be- the fresh organic matter, leaving humified organic matter (Wardle and
tween the crop and the margin in comparison to the grass margin Lavelle, 1997). Over time, the mass loss of litter in the large mesh size
(Hamza and Anderson, 2005). We used soil bulk density as a simple bags was greater than in the small mesh size bags, implying that the
surrogate to indicate the pore space, soil compaction (Buckman and activity of larger invertebrates become significant as they break down
Brady, 1960) and therefore inappropriateness of habitat and conditions this recalcitrant pool of organic matter, making it available to miner-
for soil life (Beylich et al., 2010). The greatest values of bulk density alisation (Bradford et al., 2002; Schädler and Brandl, 2005). Carrillo
were observed in the tramlines, were decomposition was the slowest. et al. (2011) observed that changes in litter decomposition processes,
Horn et al. (1995) observed that greater bulk density caused a decrease induced by the presence of meso- and macro-invertebrates, was time
in soil aeration, which in the case of the margin:crop area of this study dependent and highlighted the importance of temporal dynamics in
might indicate a slow decomposition rate and explain the observation effects on the soil fauna. The amplitude of the difference between de-
of more material remaining in the bags at the end of the experiment. composition rate in the large and small mesh sizes was the greatest in
Although it was not possible to disentangle physical losses of litter from the compacted tramline, implying that the inclusion of larger soil or-
chemical decomposition, we argue that the whole process of decom- ganisms may have supported litter decomposition despite restricted soil
position includes chemical processes as well as physical movement of conditions. However, the large mesh size bags buried in the grass
the litter (which can be caused by physical changes in soil, such as margin showed contrasted results and no difference in litter loss was
compaction, or biological transposition e.g. by earthworms). In the case observed compared to the small mesh size. This suggests that the pre-
of the tramline, we concluded that the limited losses of litter were a sence of ryegrass in the mixed litter, in an environment where microbial
consequence of compaction; whether it resulted from physical con- communities are conditioned to decompose grassy residues, would have
straints (restricted pore space resulting in limited losses of material or a synergistic effect on the decomposition of the straw from the mixture.
fauna movements), or biochemical constraints (limited oxygen supply This is a phenomenon that we could attribute to a ‘priming effect’ from
to allow efficient microbial decomposition) should be established via the ryegrass litter to amplify the decomposition of the straw in the
controlled experiments. Compaction over the whole soil profile was mixture (Fontaine et al., 2003). In the small mesh size bags, such effect
assessed by taking soil penetration resistance measurements. Penet- would therefore compensate for the effect of the soil fauna inclusion
rometer data is a measure of soil strength (Bengough et al., 2000), here observed in the large mesh size bags. Unlike the mixed litter, straw
implying that the compacted soil was stronger than that associated with residues alone decomposed faster in the large than small mesh size
crop and grass, which were not different from each other on the day the bags. This contrasting effect could result from the inabilities of the
measurements were made. Although only soil strength at the soil sur- microbial populations in the grass margin to instigate decomposition of
face was used as a data to relate to effects of compaction on litter de- wheat straw without a priming effect.
composition; all three treatments show an increase in strength with Litter quality (expressed here as C:N ratio) is well established as a
depth which is usual and probably due to the soil overburden (Horn driver of decomposition (Hamza and Anderson, 2005; Wardle and
et al., 2007). The increased strength may be due in part to differences in Lavelle, 1997) and accordingly in this study, the decomposition rate
water content and or bulk density (Bengough et al., 2000), but what- was influenced by litter type and its quality; the greater the C:N ratio of
ever the underlying causes of soil strength, the compacted soil in the the litter, the slower the decomposition. After 6 months in the soil,
tramlines was stronger than in the grass margin or in the crop (Fig. 1). significantly more litter remained in the bags containing wheat straw
All the penetrometer curves were the same basic shape apart from the than those containing ryegrass. Decomposition of mixed litter occurred
deviation at 7.4 cm depth in the tramline, which could be explained by within the interval between the wheat and ryegrass treatments; al-
wheeling pressures compacting the soil (Fig. 1). This could be inter- though the decomposition rate of the mixture was affected differently
preted as an indicator of degraded soil condition (Duiker, 2002) and by synergistic, antagonistic or additive effects of the residues depending
results in an impermeable layer of soil preventing water drainage, in- on the field area or the mesh size. It was shown by Redin et al. (2014),
creasing the likelihood that water capacity over the winter season that the diversity of functional and chemical traits of mixtures of crop

35
L. Carlesso et al. Applied Soil Ecology 134 (2019) 31–37

residues (regarding the plants alone) influences decomposition rates of and Anderson, 2005). The ban on driving on the field margin exacer-
residue mixtures. Because the effect of the mixed litter on decomposi- bates this. One simple option would be to increase the current width of
tion rates was null only in the small mesh size treatment - where only the margin supported by agri-environmental schemes from 2 to 6 m to
microbial decomposition occurred - it might be evidence for the ‘re- 12 m, and allow turning on this additional area. Grasslands are more
source concentration hypothesis’ presented by Pan et al. (2015). This resistant to compaction (Matthews et al., 2010) and we believe that
posits that the diversity of plants in a litter mixture decelerates de- such a system would minimise the “sensitive zone” and maintain soil
composition of litter because decomposers of each species suffer from a processes such as decomposition despite the high pressures and dis-
reduced availability of their preferred food resource. Because this was turbances applied in the cropped area.
not observed in the large mesh size litter bags, it implies the role of This study highlights that the current regulations for the use of grass
larger soil invertebrates regulating and promoting the microbial de- margins could be modified to optimise the ecosystem services they
composition (García-Palacios et al., 2013; Schädler and Brandl, 2005). provide. We propose that adapting the rules regarding grass margins
After 6 months decomposition, we observed that the C:N ratio of the could result in a combined benefit for growers and ecosystem services.
straw and the mixture diminished while the C:N ratio of the ryegrass For instance, extending the field margin over the compacted tramline
did not change. We posit that lower C:N ratio materials, such as rye- and allowing farmers to drive and turn in this extra-margin could result
grass, would decompose faster than high C:N ratio materials, such as in improvement of soil structure, increase in above and belowground
straw; however, we showed that changes in C:N ratio over time were biodiversity, enhancement of ecosystem services, and reduction of the
independent of decomposition rates and that the amplitude of C:N ratio costs resulting from farming this non-profitable part of the field,
changes between T0 and T6 depended on the initial C:N content. This thereby contributing to achieving more sustainable food production
change may have been driven by proximity of the litter to the soil or- systems.
ganic matter (SOM) C:N value; the closer to the SOM C:N value, the
smaller was the amplitude of change of the litter C:N value. For ex- Acknowledgements
ample, the initially high C:N value of straw had a greater amplitude of
change than the C:N value of the ryegrass, and tended towards an This work was carried out under a studentship (LC) funded by BASF.
equilibrium, similar to the average C:N ratio 11.5 found in cambisol This research was supported by Rothamsted Research (which receives
soils (Batjes, 1996). Contrary to the decomposition rate, there was a strategic funding from the UK BBSRC) and the University of
greater C:N ratio at the end of the experiment in the large mesh size Nottingham. We thank farmers Andrew & William Pitt and James
bags than in the small ones, evidence that most of the chemical de- Hinchliffe for their cooperation and for invaluable advice on arable
composition was effected by by the microbial community, while the farming systems, and Chris Watts from Rothamsted Research for his
physical breaking down of the litter was as a result of the action of advice on soil penetration resistance and use of the soil penetrometer.
larger organisms (Bradford et al., 2002). We also thank the editor and anonymous reviewers for their helpful
We have shown that prevailing soil conditions at the edge of arable suggestions to improve the manuscript.
fields affect major soil processes such as decomposition. Soil porosity is
particularly affected in this area due to heavy machinery traffic, and Appendix A. Supplementary data
inputs (fertilizers, crop residues) are less homogenously distributed
here than in the middle of the field. The uneven management and the Supplementary data to this article can be found online at https://
increased disturbance at the edge of the field are probably causal fac- doi.org/10.1016/j.apsoil.2018.10.004.
tors of the observed lower crop yields in this area. For instance, Sparkes
et al. (1998) recorded 3–19% less yield at the edge than in the middle of References
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