Geoderma 399 (2021) 115114

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Geoderma
journal homepage: www.elsevier.com/locate/geoderma

Impacts of residue quality and soil texture on soil aggregation pathways by

using rare earth oxides as tracers
S. Liu a, b, Z.C. Guo a, M. Halder a, b, H.X. Zhang a, b, J. Six c, X.H. Peng a, b, *
a
State Key Laboratory of Soil and Sustainable Agriculture, Institute of Soil Science, Chinese Academy of Sciences, Nanjing 210008, PR China
b
University of Chinese Academy of Sciences, Beijing 100081, PR China
c
Department of Environmental Systems Science, Swiss Federal Institute of Technology, ETH Zurich, CH-8092 Zurich, Switzerland

A R T I C L E I N F O A B S T R A C T

Handling Editor: Ingrid Kögel-Knabner Rare earth oxides (REOs) as tracers track aggregate formation and breakdown pathways. However, investigations
on the impacts of residue quality and soil texture on aggregation pathways are lacking. In this study, easily,
Keywords: moderately and slowly decomposable residues with different qualities (milk vetch, C/N = 16.7; maize, C/N =
Aggregate stability 35.5; and decomposed maize, C/N = 30.8) were applied to four types of red soils (Red clay, Basalt, Sandstone
Microbial activity
and Granite) with a range of soil textures. Aggregate fractions of each soil were labelled by different REOs. The
Organic residues
soil respiration (CO2), mean weight diameter (MWD) and REOs concentrations were measured after 0, 7, 14, 28
Soil aggregates
Red soils and 56 days of incubation. The results showed that except for Basalt soil, the incorporation of organic residues
increased the formation but decreased the breakdown of macroaggregates in the order of milk vetch > maize >
decomposed maize in the first two weeks, thus leading to a greater standardized MWD in higher-quality residue-
incorporated soils. In the mid-to-late incubation time (28–56 d), the standardized MWD and relative formation of
macroaggregates in the milk vetch and maize treatments converged regardless of soil types. An exponential
relation between the standardized MWD and accumulated CO2 (P < 0.05) indicated that the residue quality-
dependent MWD dynamics were further driven by microbial activity. The two coarse-textured soils (Sand­
stone and Granite) presented a more rapid response of aggregation to microbial activity than the two fine-
textured soils (Red clay and Basalt) did. The relative formation of 2–5 mm aggregates was significantly
greater in Sandstone soil at a given residue incorporation, thus leading to a faster increase in the standardized
MWD compared with fine-textured soils. However, a similar result was not observed in Granite soil. A positive
linear relationship between the standardized MWD and 2–5 mm aggregate formation in all investigated soils (P
< 0.001) suggested that MWD dynamics mainly depended on the newly formed large macroaggregate rate. Our
study demonstrates that the dynamics of soil aggregation are closely related to microbial activity and newly
formed macroaggregates, beyond the impacts of residue quality and soil texture on soil aggregation.

1. Introduction
Soil aggregation is a dynamic and complex process that includes
aggregate formation, stabilization and destabilization (Lehmann et al.,
2017; Peng et al., 2015a); it is crucial in regulating physiochemical and
biological soil properties (Six et al., 2004; Sarker et al., 2018), moti­
vating soil microbial activities (Zhang et al., 2018) and influencing soil
carbon and nutrient cycling (Bosshard et al., 2008; Nesper et al., 2015;
Rahman et al., 2019). However, soil aggregate dynamics have been
elusive because of limited tracking methods. Studies have reported that
rare earth oxides (REOs) can be uniformly incorporated into soil
aggregates of different sizes (Zhang et al., 2001; Stevens and Quinton,
2008) because of their strong binding affinity with mineral surfaces
within soil (Kimoto et al., 2006), which makes it possible to track the
formation and breakdown of soil aggregates. The REO approach was
proposed by Zhang et al. (2001) to explore soil aggregation in a soil
column study and subsequently improved by De Gryze et al. (2006) and
Peng et al. (2017). Previous studies on Alfisols (De Gryze et al., 2006;
Morris et al., 2019), Ultisols (Peng et al., 2017) and Vertisols (Rahman
et al., 2019) have proven that the approach of using REOs as tracers is
universal for tracking aggregate dynamics (Liu et al., 2019).
By using REOs and isotopically labelled carbon as tracers, Peng et al.

* Corresponding author at: State Key Laboratory of Soil and Sustainable Agriculture, Institute of Soil Science, Chinese Academy of Sciences, Nanjing 210008, PR
China.
E-mail address: xhpeng@issas.ac.cn (X.H. Peng).

https://doi.org/10.1016/j.geoderma.2021.115114
Received 25 August 2020; Received in revised form 24 March 2021; Accepted 24 March 2021
Available online 4 May 2021
0016-7061/© 2021 Elsevier B.V. All rights reserved.
S. Liu et al. Geoderma 399 (2021) 115114

Table 1
Description of soil sampling site, annual precipitation, parent material, taxonomic classification, clay minerals and land-use.
Soil types Sampling sites Annual precipitation (mm) Parent materials USDA Soil taxonomy Clay minerals Land use

Red clay Yingtan, Jiangxi 1750 Quaternary red clay Typic Plinthudult Ultisol Kaolinite, haematite Peanut field
Sandstone Yingtan, Jiangxi 1750 Sandstone Typic Hapludult Ultisol Kaolinite, halloysite Grassland
Basalt Danzhou, Hainan 1815 Basalt Typic Hapludox Oxisol Laolinite, hydromica Banana garden
Granite Xingguo, Jiangxi 1515 Granite Typic Paleudult Ultisol Kaolinite, vermiculite, chlorite Grassland

(2017) determined soil aggregate transformation paths following the
input of organic matter (glucose) in red clay soil and proposed a first-
order kinetic model for aggregate dynamics. Rahman et al. (2019)
investigated the effects of residue quality (maize straw, C/N = 32;
miscanthus straw, C/N = 220) and nitrogen input on soil aggregate
turnover in Shajiang black soil and concluded that aggregate turnover in
the soil incorporated with maize straw was faster than that in the soil
incorporated with miscanthus straw. Morris et al. (2019) studied how
aggregate turnover times were affected by arbuscular mycorrhizal fungi
(AMF) in an Albic Luvisol and showed that AMF increased the large
macroaggregate formation and slowed the disintegration of large and
small macroaggregates. These studies both focused on soil aggregate
turnover time and formation or breakdown processes in one type of soil
with no more than two organic residue applications, but less attention
has been given to the effects of residue quality, soil texture and their
interactions on soil aggregate dynamics.
Straw residue amendment significantly enhanced microbial biomass
carbon (MBC) and phospholipid fatty acids (PLFAs) (Rahman et al.,
2017), thereby increasing the formation of macroaggregates and
consequently affecting aggregate stability (Chivenge et al., 2011; Sall
et al., 2016; Rahman et al., 2019). However, the effects of residue
quality on soil aggregation are inconsistent in terms of the C/N ratio.
Based on an eight-month mesocosm experiment in red clay soil, Chiv­
enge et al. (2011) found that relative to high-quality organic residues,
low-quality organic residues resulted in greater aggregate stability in red
clay soil. From a 120-d incubation experiment in a loamy-sand soil, Sall
et al. (2016) reported that the formation of macroaggregates and
aggregate stability was not significantly affected by the quality of resi­
dues in loamy sand soil. Rahman et al. (2019) suggested that relative to
poor-quality organic residues, higher decomposition of high-quality
organic residues resulted in better soil aggregation in a Vertisol during
28-day incubation time. These inconsistent results may be dependent on
the phase of residue decomposition. According to Monnier’s conceptual
model (Monnier, 1965), easily decomposed residues may have a
considerable improvement on aggregation in the short term, but the
positive effect will be attenuated with increasing time, whereas this
trend is the opposite for slowly decomposed residues (Abiven et al.,
2008). Soil aggregate formation and breakdown processes are the inner
drivers for the dynamics of the MWD. However, how the MWD changes
with soil aggregate formation and breakdown processes under different
phases of residue decomposition remains unclear.
In addition to residue quality, residue decomposition was reportedly
affected by soil texture, but contrasting results were observed. The
decomposition rates of organic residues were reported to be faster in
coarse-textured soil than in fine-textured soil (Parwada and Van Tol,
2018; He et al., 2020), while the opposite result was found by
Hamarashid et al. (2010). Soil texture is another key factor in soil ag­
gregation (Bronick and Lal, 2005; Wick et al., 2009). A high correlation
between clay content and aggregate stability was observed by previous
investigations (Adhikari and Bhattacharyya, 2015; Blanco-Moure et al.,
2016; Rivera and Bonilla, 2020). Clay particles favour increasingly
stronger organic C-clay interactions due to their larger specific surface
areas, thus producing a good soil structure (von Lützow et al. 2006).
However, different results of soil aggregate formation have been
observed. Denef and Six (2005) reported that residue input and plant
growth had a greater positive effect on macroaggregate formation in
illitic (clay = 27%) than in kaolinitic soil (clay = 85%). In contrast,
Gentile et al. (2011) found that residue additions increased macro­
aggregation in clayey soil more than in loamy sand soil. Such inconsis­
tent results call for more detailed information on how soil aggregate
formation and breakdown processes vary with soil texture.
In this study, rare earth oxides (REOs) were used as tracers to track
aggregation pathways. Easily, moderately and slowly decomposable
residues with different qualities (milk vetch, C/N = 16.7; maize straw,
C/N = 35.5; and decomposed maize straw, C/N = 30.8) were applied to
four types of red soils developed from different parent materials (Red
clay, Basalt, Sandstone and Granite) with a range of soil textures. Our
objectives were 1) to determine the impacts of residue quality, soil
texture and their interactions on soil aggregate formation and break­
down pathways and 2) to analyze how soil aggregate formation and
breakdown processes control soil aggregation dynamics during residue
decomposition.
2. Materials and methods
2.1. Soil sampling and properties
Four red soils derived from different parent materials were collected
from subtropical and tropical China, including Red clay, Sandstone,
Granite and Basalt (Table 1). All soil samples were collected from top
layer (0–10 cm) and air-dried at room temperature, sieved to < 2 mm,
with fine roots and other plant debris carefully removed. Soil properties
were determined by routine methods (Lu, 1999). Soil pH was measured
on suspension formed from a soil: deionized water ratio of 1:2.5 by a pH
meter (PHSJ-4F, Shanghai Electronics Science Instrument Co.,
Shanghai, China). The contents of soil organic carbon (SOC) and total
nitrogen (TN) were determined with an elemental analyzer (vario MAX
CN, Elementar, Germany). CEC was measured by the ammonium acetate
method. A more detailed description of the Fe/Al oxides determination
was presented in Peng et al. (2015b). Clay mineralogy was qualified
using X-ray diffraction analysis and additional information was obtained
by scanning electron microscopy. Soil texture was determined by the

Table 2
Selected physical and chemical properties of the soils studied.
Soil types Field capacity pH SOC† TN Fe2O3 Al2O3 Sand Silt Clay C/N CEC
− 1 − 1 1
gg g kg cmol kg−

Red clay 0.4 4.71 7.66 0.73 59.0 166.1 281.2 337.2 381.6 10.5 9.4
Sandstone 0.22 4.85 3.07 0.31 19.5 66.6 754 136.8 109.2 9.9 4.8
Basalt 0.4 6.54 6.53 0.58 16.5 251.5 262 362.8 375.2 11.3 8.3
Granite 0.24 4.86 8.24 0.77 47.8 154.0 600.4 287.6 112 10.7 8.9

Note: SOC, soil organic carbon; TN, total nitrogen; MWD, mean weight diameter. Sand (>0.05 mm), silt (0.05–0.002 mm), clay (<0.002 mm).

2
S. Liu et al.
pipette method. Field capacity was the gravimetric moisture value at −
33 kPa. Soil properties were listed in Table 2.
2.2. Preparation of REO-labelled soils
To capture the inherent structure of soil, natural aggregates
following the wet labelling method is recommended in this study (Liu
et al., 2019). Four portions of wet-labelled soils were obtained after
mixing four REOs (La2O3, Sm2O3, Nd2O3 and Gd2O3) with < 2 mm soil
separately. Briefly, each REO powder was suspended in deionized water
by vortex mixing, then sprayed gradually into air-dry soil to achieve a
concentration of 500 mg REO kg− 1 soil. The soil was gently mixed while
spraying by rotating it. After mixing and adjusting to 60% of field ca­
pacity water content, the REO-labelled soils were kept at 4 ℃ for 7 days
to allow water equilibration with minimal microbial activity.
The soils were oven-dried at 40 ℃ for two days and sieved to < 5
mm. Then wet-sieving was conducted to obtain four aggregate fractions
following the Elliot’s method (1986): (i) large macroaggregates (2–5
mm); (ii) small macroaggregates (0.25–2 mm); (iii) microaggregates
(0.053–0.25 mm); (iv) silt and clay sized aggregates (<0.053 mm),
indicated A, B, C and D fractions in this study, respectively. The frac­
tioned aggregates were oven-dried at 40 ℃ and weighed. Aggregates
from A-D fractions were recombined into a new soil sample in which
each aggregate fraction contained a different REO tracer: A fraction
labelled by Nd2O3, B fraction labelled by Sm2O3, C fraction labelled by
Gd2O3, and D fraction labelled by La2O3 (Fig. S1, Supplementary
materials).
2.3. Experimental design
In order to evaluate the impact of residue quality on soil aggregation
pathways, easily, moderately and slowly decomposable residues were
involved in this incubation study: milk vetch (Astragalus sinicus L.) (C =
31.1%, C/N = 16.7), maize straw (Zea mays L.) (C = 37.6%, C/N = 35.5)
and decomposed maize straw (C = 42.2%, C/N = 30.8). To obtain the
decomposed maize straw, 350 g maize straw was added with 35 ml
diluted straw decomposition maturing agent (8.2 × 108/ml), and then
decomposed at 30 ℃ for 20 days at 60% of saturation moisture content
(371 g water kg− 1 maize straw) (Li et al., 2015). Thereafter, the
decomposed maize straw was oven-dried at 40 ◦ C, and then sterilized at
121 ◦ C for 2 h in the autoclave, which repeated three times (Ni and Pan,
2018). Milk vetch straw, maize straw and decomposed maize straw were
crushed to < 0.25 mm for the use of the following incubation
experiment.
For each type of soil, four treatments were designed as follows: i)
REO labelled soil without any straw as a control (Control), ii) REO
labelled soil added with milk vetch (Vetch), iii) REO labelled soil added
with maize straw (Maize), and iv) REO labelled soil added with
decomposed maize straw (Decomposed maize). Each treatment was
triplicate. The amount of 1% on dry weight has been applied in a set of
previous investigations (Liu et al., 2019; Rahman et al., 2019), hence
these three types of residues were incorporated into the REO-labelled
soils by the same amount. Straw was mixed homogeneously with the
REO-labelled soil in plastic packaging bag to avoid aggregates break­
down. To maintain a uniform condition, the soil of the control treatment
was subjected to the same procedure.
The soil-straw mixture was gently packed into a PVC core (4 cm
diameter, 5 cm height) step by step with a flat presser to get a bulk
density of 1.2 g cm− 3, followed by wetting to 60% of the field capacity.
There were 240 soil cores in total, which were divided equally into 5 big
boxes. Each box contained 48 samples. All soil columns were kept in the
incubator at 4 ◦ C for 3 days to allow the water balance with minimal soil
microbial activities and then transferred to an incubation chamber at 25
± 2 ◦ C. Soil cores were then placed in 500 ml glass jar and incubated in
plastic wrap sealed condition for 56 days. 20 holes were made by nee­
dles on each warp to allow air exchange. Soil water content was kept
3
Geoderma 399 (2021) 115114
controlled during the entire incubation experiment by regularly adding
deionized water to soil columns.
Aggregate dynamics were measured by destructively harvesting
batches of soil on days 0, 7, 14, 28 and 56 of incubation. For each
sampling day, we collected the entire soil cores for further analysis from
four treatments, 3 replicates for every treatment, hence 48 soil cores (4
× 3 × 4) were collected from four types of soils. Collected soil cores were
oven-dried at 40 ◦ C for 48 h and broken down to < 5 mm before
measuring the aggregate size distribution and REOs contents. Soil
respiration was analyzed on days 0, 1, 3, 5, 7, 11, 14, 28, 42 and 56 of
incubation.
2.4. Soil measurements
2.4.1. Soil respiration
As detailedly described by Rahman et al., (2019), ventilated and
sealed glass jars (500 ml) were used for CO2 fluxes determination. For
each gas collecting, two 20 ml glass vials were collected. One for CO2
concentration analyses and one for backup. The second gas collection
was taken after 6 h of closure. After that, the glass jars were covered with
polyethylene film and incubated for the next sampling round. The CO2
concentrations were analyzed on a gas chromatograph (GC) (Agilent
7890, Santa Clara, USA) equipped with a thermal conductivity detector
(analytical precision ± 0.5%).
2.4.2. Aggregate separation
Separation of aggregates was done by the wet sieving method
(Elliott, 1986). Briefly, the soil was submerged for 5 min in deionized
water, then a series of sieves were used to obtain four aggregate size
fractions: >2 mm (large macroaggregates), 0.25–2 mm (small macro­
aggregates), 0.053–0.25 mm (microaggregates) and < 0.053 mm (silt
and clay fractions). Aggregates were separated by slowly moving the
sieve up and down to about 3 cm depth in water for 50 times over 2
mins. The aggregates remaining on each sieve were oven-dried at 40 ◦ C
for 48 h and weighed. Then the mean weight diameter (MWD) of ag­
gregates was calculated as follows:
∑
n
MWD = xi × wi (1)
i=1
where xi is the mean diameter of each aggregate fraction, wi is the
mass proportion of aggregate fraction remaining on each sieve, and n is
the number of fractions.
In order to remove the effect of initial conditions, standardized MWD
was proposed as following:
MWDt
StandardizedMWDt = (2)
MWD0
where MWDt is the mean weight diameter on t sampling day (t = 0, 7,
14, 28 or 56 d), MWD0 is the initial MWD on 0d during the incubation
period. The standardized MWD on 0 d is calculated as 1 in all treatments.
2.4.3. REOs concentrations
The contents of each REO were measured by ICP-MS after alkaline
fusion (Bayon et al., 2009; Peng et al., 2017). Briefly, 0.125 g of soil was
placed into a corundum crucible, mixed with 1 g Na2O2 and covered by
0.25 g Na2O2. The crucible was put in a muffle furnace at 700 ◦ C for 15
min. To remove the resulting H2O2 and complete co-precipitation, the
beaker contained crucible with 100 ml distilled water was heated to
200 ◦ C on a hotplate for 3 h. The precipitate was rinsed into a 250 ml
volumetric flask to which 2 drops of phenolphthalein indicator was
added and then titrated by 1% HNO3 to the color change point. A further
2 ml HCl was added to the volumetric flask to maintain acidity. Then 15
ml sample solution was transferred into an HDPE bottle and analyzed for
REOs by high resolution inductively coupled plasma mass spectrometry
(HR-ICP-MS: Finnigan Element, USA).
S. Liu et al. Geoderma 399 (2021) 115114

B fraction labelled by Sm2O3, C fraction labelled by Gd2O3, and D

fraction labelled by La2O3. There are 12 possible pathways in total,
including 6 potential breakdown pathways into smaller aggregate
fractions (a-f) and another 6 (g-l) reformation pathways into larger
aggregate fractions (Fig. 1). For example, a indicates the pathway of A
fraction breakdown into B fraction, whereas g indicates the pathway of B
fraction buildup into A fraction.
The transfer portions among soil aggregates can be calculated based
on the changes of REOs concentrations. These transfers between time t1
and time t2 can be summarized in a discrete transformation matrix K(t2-
t1):
⎡ ⎤
1− a− d− f g j l
⎢ a 1− g− b− e h k ⎥
K(t2 − t1 ) = ⎣ ⎦
d b 1− j− h− c i
f e c 1− l− k− i
(3)
Fig. 1. The 12 possible pathways between four different aggregate fractions (a-
where the K(t2-t1) indicates the change in the proportions of aggre­
f are breakdown direction and g-l are build-up direction). A, B, C and D indicate
2–5 mm, 0.25–2 mm, 0.053–0.25 mm and < 0.053 mm, respectively. gates falling into sizes A, B, C or D between time steps t1 and t2.
In the aggregate breakdown (BD) or build-up (BU) direction, the
changes in aggregate proportion of A, B and C fractions during the in­
2.5. Theory of aggregation pathways
cubation from time t1 to t2 are expressed as follows:
The mathematical calculation of aggregate turnover by REO tracers BD(A) = (at2 − at1 ) + (dt2 − dt1 ) + (ft2 − ft1 ) (4)
was described by Peng et al. (2017). Briefly, aggregates from A-D frac­
tions were tracked by different REO tracer: A fraction labelled by Nd2O3, BD(B) = (bt2 − bt1 ) + (et2 − et1 ) (5)

Control Vetch Maize Decomposed maize

250 Red clay 1200 Red clay
200 900
150
600
100
50 300
0 0
Soil respiration CO2 mg kg-1 soil day-1

Accumulated respirition CO2 mg kg-1 soil

Basalt Basalt
250 1200
200
900
150
600
100
50 300
0 0

Sandstone Sandstone
250 1200
200 900
150
600
100
50 300

0 0

250 Granite 1200

Granite
200 900
150
600
100
50 300
0 0
0 10 20 30 0 10 20 30

Incubation time (d)

Fig. 2. The daily (left) and accumulated (right) soil respiration in the first four weeks incubation under treatments of i) REO labelled soil without any straw as a
control (Control), ii) REO labelled soil added with milk vetch (Vetch), iii) REO labelled soil added with maize straw (Maize), and iv) REO labelled soil added with
decomposed maize straw (Decomposed maize).

4
S. Liu et al. Geoderma 399 (2021) 115114

Control Vetch Maize Decomposed maize

8 Red caly 8 Sandstone
7 7
6 6
5 5
4 4
Standardized MWD 3 3
2 2
1 1
0 0
8 Basalt 8 Granite
7 7
6 6
5 5
4 4
3 3
2 2
1 1
0 0
0 7 14 28 56 0 7 14 28 56

Incubation time (d)

Fig. 3. The dynamics of relative aggregate stability (standardized MWD) over time under i) REO labelled soil without any straw as a control (Control), ii) REO
labelled soil added with milk vetch (Vetch), iii) REO labelled soil added with maize straw (Maize), and iv) REO labelled soil added with decomposed maize straw
(Decomposed maize).

BD(C) = (ct2 − ct1 )
(gt2 − gt1 )×mB + (jt2 − jt1 ) × mC + (lt2 − lt1 ) × mD
BU(A) =
mA
(ht2 − ht1 ) × mC + (kt2 − kt1 ) × mD
BU(B) =
mB
(it2 − it1 )×mD
BU(C) =
mC
where mA , mB , mC and mD are the initial mass of aggregate fraction A,
B, C and D, respectively. In breakdown direction, a negative value (BD <
0) means less aggregate disruption relative to the initial condition,
whereas a positive value means (BD > 0) a greater disruption. In the
build-up direction, a positive value means (BU > 0) more aggregate
build-up relative to the initial condition, whereas a negative value (BU
< 0) means aggregate breakdown rate.
2.6. Statistical analysis
Analysis of variance (ANOVA) was performed at each sampling date
to test the effects of residue quality and soil texture on aggregate size
distribution, mean weight diameter (MWD), soil respiration and relative
change in formation or breakdown direction (SPSS, 2004). The least
significant difference (LSD at P < 0.05) test was applied to assess the
differences among the means of three replicates (n = 3).
3. Results
3.1. Soil respiration
During the late stage of incubation (28–56 d), the respiration rates of
soil in the milk vetch, maize and decomposed maize treatments were
similar but low; hence, the soil respiration in the first four weeks was
(6) displayed in Fig. 2. The incorporation of organic residues resulted in
greater soil respiration relative to the control treatment in all investi­
gated soil types (P < 0.05). A pronounced peak in CO2 occurred on the
(7) first day and decreased rapidly over the following days during the entire
incubation period. During the initial 14 days of incubation, the highest
soil respiration rate was observed with milk vetch addition, followed by
(8) maize and then decomposed maize. The cumulative respiration signifi­
cantly differed among treatments in the four types of soils (P < 0.01,
Fig. 2). The cumulative respiration was in the order of milk vetch >
(9)
maize > decomposed maize > control treatment. The soil respiration
rate and accumulative respiration during the 56-day incubation were
lower in Granite soil than in the other three soils (e.g., 365 vs 888–1082
mg kg− 1 soil with milk vetch addition).
3.2. Soil aggregate stability
The aggregate size distributions in the four soils over the incubation
time were displayed in Figs. S2–S5 (supplementary material). The
changing patterns of relative aggregate stability, indicated by the stan­
dardized MWD and depended on soil texture and residue quality, were
shown in Fig. 3. In the control treatment, the standardized MWD
remained stable over time in all soils. In the first 14-day incubation time,
the standardized MWD was enhanced in the order of milk vetch > maize
> decomposed maize relative to the control in Red clay, Sandstone and
Granite soils, whereas in Basalt soil, the standardized MWD with maize
application was the highest. In the late period of incubation (28–56
days), the standardized MWDs in the milk vetch and maize treatments
converged. In Red clay soil, the standardized MWD increased consid­
erably in the first two weeks and then sharply decreased afterwards with
milk vetch and maize additions, while the highest standardized MWD in
the decomposed maize treatment was observed on day 28. The pro­
nounced peak occurred on day 7 in Sandstone soil under milk vetch
treatment; the standardized MWD increased during the initial two-week
incubation and remained constant for the rest of the incubation time
under maize and decomposed maize treatments. Instead, the

5
S. Liu et al. Geoderma 399 (2021) 115114

Red clay Sandstone

8 8 -0.003x
y=1.54(1-e-0.005x)+1 y=5.99(1-e )+1
R2=0.43* 2 **
R =0.88
6 6

4 4

Standardized MWD
2 2

0 0
0 200 400 600 800 1000 0 200 400 600 800 1000
8 8
Basalt Granite
y=1.23(1-e-0.002x)+1 y=1.50(1-e-0.012x)+1
2 * 2 **
6 R =0.63 6 R =0.68

4 4

2 2

0 0
0 200 400 600 800 1000 1200 0 200 400

Accumulated respiration (CO2)(mg kg-1 soil)

Fig. 4. An exponential relationship between relative aggregate stability (standardized MWD) with residue straw addition and accumulated soil respiration.

Fig. 5. The 12 transformation paths of four aggregate fractions on days 0, 7, 14, 28 and 56 of incubation in Red clay soil under four treatments of i) REO labelled soil
without any straw as a control (Control), ii) REO labelled soil added with milk vetch (Vetch), iii) REO labelled soil added with maize straw (Maize), and iv) REO
labelled soil added with decomposed maize straw (Decomposed maize). Values in arrows are the relative change of this aggregate fraction (%). A, B, C, and D
represent 2–5 mm, 0.25–2 mm, 0.053–0.25 mm, and < 0.053 mm aggregates, respectively.

6
S. Liu et al. Geoderma 399 (2021) 115114

Fig. 6. The 12 transformation paths of four aggregate fractions on days 0, 7, 14, 28 and 56 of incubation in Basalt soil under four treatments of i) REO labelled soil
without any straw as a control (Control), ii) REO labelled soil added with milk vetch (Vetch), iii) REO labelled soil added with maize straw (Maize), and iv) REO
labelled soil added with decomposed maize straw (Decomposed maize). Values in arrows are the relative change of this aggregate fraction (%). A, B, C, and D
represent 2–5 mm, 0.25–2 mm, 0.053–0.25 mm, and < 0.053 mm aggregates, respectively.

Fig. 7. The 12 transformation paths of four aggregate fractions on days 0, 7, 14, 28 and 56 of incubation in Sandstone soil under four treatments of i) REO labelled
soil without any straw as a control (Control), ii) REO labelled soil added with milk vetch (Vetch), iii) REO labelled soil added with maize straw (Maize), and iv) REO
labelled soil added with decomposed maize straw (Decomposed maize). Values in arrows are the relative change of this aggregate fraction (%). A, B, C, and D
represent 2–5 mm, 0.25–2 mm, 0.053–0.25 mm, and < 0.053 mm aggregates, respectively.

7
S. Liu et al.
Fig. 8. The 12 transformation paths of four aggregate fractions on days 0, 7, 14, 28 and 56 of incubation in Granite soil under four treatments of i) REO labelled soil
without any straw as a control (Control), ii) REO labelled soil added with milk vetch (Vetch), iii) REO labelled soil added with maize straw (Maize), and iv) REO
labelled soil added with decomposed maize straw (Decomposed maize). Values in arrows are the relative change of this aggregate fraction (%). A, B, C, and D
represent 2–5 mm, 0.25–2 mm, 0.053–0.25 mm, and < 0.053 mm aggregates, respectively.
standardized MWD presented a continuously increasing trend with time
regardless of residue quality in Basalt soil, with the order of maize >
milk vetch > decomposed maize. Similar to the standardized MWD
dynamics in Basalt soil, increasing patterns were observed with maize
and decomposed maize applications in Granite soil.
With residue incorporation, the standardized MWD was exponen­
tially associated with accumulated respiration during the incubation
period regardless of soil type, showing that the MWD dynamics were
further driven by microbial activity (P < 0.05, Fig. 4). After taking the
derivative of these fitting formulas, the changing rates of the standard­
ized MWD of Red clay, Basalt, Sandstone and Granite soils were 0.008,
0.002 and 0.018, 0.018, respectively, indicating that the MWD respon­
ded more rapidly to soil microbial activity in coarse- than fine-textured
soil. At a given residue incorporation, the standardized MWD increased
faster in Sandstone soil than in the other three types of soils (P < 0.05,
Fig. S6, supplementary material).
3.3. Aggregate transformation paths
The dynamics of soil aggregation are controlled by aggregate trans­
formation. Aggregate transformation paths were calculated based on
changes in REOs concentrations among four aggregate size fractions
over time (Figs. 5–8). On day 0, the recombination and wet sieving effect
resulted in changes in soil aggregates. Relative to day 0, the differences
in transformation paths on days 7, 14, 28 and 56 were caused by the
incubation effect.
Except for Basalt soil, the addition of organic residues significantly
reduced aggregate breakdown (P < 0.001) and increased aggregate
formation (P < 0.001) in the order of milk vetch > maize > decomposed
8
Geoderma 399 (2021) 115114
maize treatments in the first two weeks compared with the control
treatment (Figs. 5–8). Taking day 7 as an example, relative to the control
treatment, the transformation portions of A aggregates in the breakdown
direction with milk vetch, maize and decomposed maize applications
were reduced by 36, 16 and 6 percentage points, respectively (marked
with red boxes in Fig. 5). On the other hand, the transformation portions
of A aggregates in the build-up direction with milk vetch, maize and
decomposed maize applications were increased by 55, 32 and 10 per­
centage points, respectively, on day 7 of incubation (marked in red bold
font in Fig. 5).
A greater portion of aggregates transformed between neighbouring
size fractions in all treatments either in the breakdown or build-up di­
rection, except the 2–5 mm aggregates in the breakdown direction in all
investigated soils (Figs. 5–8). Taking the Red clay soil on day 7 as an
example, the transformation portions of B → C and B → D were 63% and
18% under the control treatment and 47% and 9% under the milk vetch
treatment (marked with green boxes in Fig. 5); the transformation
portions of D → C and D → B were 54% and 21% under the control
treatment and 27% and 17% under the milk vetch treatment (marked
with orange circles in Fig. 5). The same patterns were observed with the
additions of maize and decomposed maize. In the four types of soils, the
changes in the build-up directions mainly occurred in the first two weeks
and then attenuated with time with or without residues (Figs. 5–8),
which is in agreement with the dynamics of the standardized MWD and
daily CO2 over time. For example, the transformation portion of A ag­
gregates in the build-up direction increased from 55% on day 7 to 73%
on day 14 and then decreased to 21% at the end of incubation under the
milk vetch treatment in Red clay soil (marked with diagonal boxes in
Fig. 5).
S. Liu et al. Geoderma 399 (2021) 115114

Red clay Basalt Sandstone Granite

40 y=0.18+1.01x 40 y=-0.01+0.99x 50 y=0.50+0.97x 40 y=0.35+0.97x
2-5 mm R2=1.00*** R2=0.97*** 2
40 R =1.00*** R2=0.98***
30 30 30
30
20 20 20
20
10 10 10 10

0 0 0 0
Predicted aggregates (%)

0 10 20 30 40 0 10 20 30 40 0 10 20 30 40 50 0 10 20 30 40

30 y=--0.61+1.03x 60 y=0.31+0.97x 20 y=--0.07+1.09x 50 y=0.23+0.99x

0.25-2 mm

R2=0.95** R2=0.95** R2=0.72** 40 R2=0.82**

20 40
30
10
20
10 20
10
0 0 0 0
0 10 20 30 0 20 40 60 0 10 20 0 10 20 30 40 50
0.053-0.25 mm

40 y=1.11+1.06x 30 y=-1.21+1.03x 80 y=-2.93+1.00x 70 y=1.79+0.93x

R2=0.93** R2=0.90** 2 60 R2=0.87**
30 60 R =0.94** 50
20
40
20 40
30
10 20
10 20
10
0 0 0 0
0 5 10 15 20 25 30 35 40 0 10 20 30 0 20 40 60 80 0 10 20 30 40 50 60 70

70 y=0.08+0.96x 40 y=1.45+1.01x 30 y=5.81+0.82x 30 y=3.90+0.90x

60 R2=0.97**
<0.053 mm

R2=0.98** R2=0.68** R2=0.73**

50 30
20 20
40
20
30
20 10 10
10
10
0 0 0 0
0 10 20 30 40 50 60 70 0 10 20 30 40 0 10 20 30 0 10 20 30

Measured aggregates (%)

Fig. 9. The relationship between measured and predicted aggregates.

As in two fine-textured soils (Red clay and Basalt), the build-up
transformation portion of 2–5 mm was significantly smaller than that
in two coarse-textured soils (Sandstone and Granite) in the first two
weeks with milk vetch application, while the opposite pattern was
observed in the maize and decomposed maize treatments after 14 days
of incubation (P < 0.05) (Figs. 5–8). With milk vetch addition, taking
day 7 as an example, the transformation proportions of B → A in Red
clay and Basalt soils were 20% and 14%, respectively, which were
remarkably smaller than those in Sandstone and Granite soils (57% and
37%).
As reported by Peng et al. (2017), the soil aggregate proportions can
be predicted by the sum of the transformation portions in the build-up
and breakdown directions. The relationship between the measured
and predicted aggregates for the four fractions was significantly linear
(P < 0.01, Fig. 9). Relative to coarse-textured soils, the REO labelling
method worked better in fine-textured soils.
3.4. Relative change in the breakdown and build-up directions
The relative changes in aggregate fractions in the breakdown or
build-up direction were displayed in Figs. 10 and 11, respectively. In the
breakdown direction, a negative value means less aggregate disruption
relative to the initial condition, whereas a positive value means a greater
disruption (Fig. 10). Compared with the control treatment, the incor­
poration of all three organic residues presented more negative values,
indicating less aggregate disruption in all investigated soils. Only
0.053–0.25 mm aggregates in Basalt soil and 0.25–2 mm aggregates in
Granite soil presented positive values but were still smaller than those of
the control treatment. The breakdown of soil aggregates decreased with
time in the first two weeks and accelerated afterwards, except for 2–5
mm aggregates in Granite soil and 0.25–2 mm aggregates in Sandstone
soil with maize application, which showed a decreasing trend over the
whole incubation period. In the first two weeks, the reduction in
aggregate breakdown was generally in the order of milk vetch > maize
> decomposed maize in Red clay, Sandstone and Granite soils, while this
reduction was the highest with maize straw addition, followed by milk
vetch and decomposed maize in > 0.25 mm aggregates in Basalt soil.
In the build-up direction, a positive value indicates more aggregate
build-up relative to initial conditions, whereas a negative value indicates
an aggregate breakdown rate (Fig. 11). The incorporation of three
organic residues had greater values than the control treatment for >
0.25 mm aggregates, but the opposite was true for 0.053–0.25 mm ag­
gregates. In the first two weeks, the relative formation of 2–5 mm ag­
gregates was in the order of milk vetch > maize > decomposed maize >
control in Red clay, Sandstone and Granite soils, whereas the highest
value was found in the maize treatment in Basalt soil. Except for Granite

9
S. Liu et al. Geoderma 399 (2021) 115114

Control Vetch Maize Decompsed maize

2-5 mm 0.25-2 mm 0.053-0.25 mm
0.6 0.6 0.6
Red clay

0.0 0.0 0.0

-0.6 -0.6 -0.6

Relative change in aggregates (%)

0.6 0.6 0.6

Basalt

0.0 0.0 0.0

-0.6 -0.6 -0.6

0.6 0.6 0.6

Sandstone

0.0 0.0 0.0

-0.6 -0.6 -0.6

0.6 0.6 0.6

Granite

0.0 0.0 0.0

-0.6 -0.6 -0.6

0 7 14 28 56 0 7 14 28 56 0 7 14 28 56

Incubation time(d)
Fig. 10. Relative changes in aggregates in the breakdown direction.

soil, the relative formation of soil aggregates in the milk vetch and maize
treatments converged in the late period of incubation (28–56 days). In
Red clay soil, the relative formation of 2–5 mm aggregates increased to
28 d of incubation and then decreased over time. The relative formation
of 2–5 mm aggregates accelerated in the first 14 days and remained
constant in the mid-to-late incubation time (28–56 d) in Sandstone soil,
while an increasing trend was observed during the entire incubation
time in Basalt and Granite soils. Compared with 2–5 mm aggregates, the
relative formation of 0.053–0.25 mm aggregates was negligible. A
significantly linear relationship between the standardized MWD and
relative formation in 2–5 mm aggregates was observed in all soils (R2 >
0.94, P < 0.001, Fig. 12), suggesting that the dynamics of the MWD
mainly depended on newly formed large macroaggregates in this study.
4. Discussion
The impact of residue quality on soil aggregation depends on the
decomposition phase, varying from the early (0–14 d) to mid-to-late
(28–56 d) stage of incubation (Fig. 3). Except for Basalt soil, the stan­
dardized MWD was in the order of milk vetch > maize > decomposed
maize in the first two weeks, while the values with milk vetch and maize
addition converged in the late period of incubation (Fig. 3). Similar
results were reported by Rahman et al. (2019) who found that higher
formation and lower breakdown of aggregates resulted in a significantly
higher MWD with high-quality maize straw (C/N = 32) relative to low-
quality miscanthus straw (C/N = 220) over time. Following Monnier’s
conceptual model (Abiven et al., 2008), milk vetch, maize and decom­
posed maize straws were regarded as easily, moderately and slowly
decomposable residues, respectively. Milk vetch straw, which is rich in
labile polysaccharides relative to maize and decomposed maize, acts as a
C source for microbes in residue decomposition (Sarker et al., 2018;
Halder et al., 2021). The resultant microbial products facilitate aggre­
gate formation (Alami et al., 2000), thus resulting in an initial sharp
increase in the MWD. Maize straw is rich in O-alkyl C and di-O-alkyl C
fractions and mainly associated with carbohydrates such as cellulose,
positively influenced the aggregate stability mid-to-long-term persis­
tence (Abiven et al., 2007; Sarker et al., 2018; Halder et al., 2021). The
low decomposition rate in decomposed residues (Fig. 2) provides a
continuous release of nutrients capable of sustaining microbial pop­
ulations for long time periods (Murphy et al., 2007); hence, a slight but
progressive increase in the MWD is observed (Sarker et al., 2018).
Majumder and Kuzyakov (2010) reported that in the first phase (0–15
days) of residue decomposition, non-cellulosic polysaccharides, proteins
and hemicellulose, which were readily available to microorganisms,

10
S. Liu et al. Geoderma 399 (2021) 115114

Control Vetch Maize Decomposed maize

2-5 mm 0.25-2 mm 0.053-0.25 mm
80 1.2 1.2
70
Red clay 60
50 0.6 0.6
40
30
20 0.0 0.0
10
0
-10 -0.6 -0.6
Relatvive change in aggregates (%)

70 1.2 1.2
60
50
0.6 0.6
Basalt

40
30
20 0.0 0.0
10
0
-10 -0.6 -0.6

140 1.2 1.2

120
Sandstone

100
80 0.6 0.6
60
40 0.0 0.0
20
0
-20 -0.6 -0.6

12 1.2 1.2
10
8 0.6 0.6
Granite

6
4
2 0.0 0.0
0
-2 -0.6 -0.6
0 7 14 28 56 0 7 14 28 56 0 7 14 28 56

Incubation time (d)

Fig. 11. Relative changes in aggregates in the build-up direction.

5 Red clay 9 Sandstone

y=-22.0+22.4x 8 y=-17.35+17.79x
4 R2=0.96*** 7 R2=0.97***
6
3
5
4
2
3
Standardized MWD

1 2
1
0 0
-10 0 10 20 30 40 50 60 70 80 90 0 20 40 60 80 100 120 140
3 Basalt 4 Granite
y=-36.15+38.54x y=-3.88+4.24x
R2=0.94*** 3 R2=0.97***
2

1
1

0 0
-10 0 10 20 30 40 50 60 70 -2 0 2 4 6 8 10 12 14

Relative change in 2-5 mm aggregate in the build up direction (%)

Fig. 12. Relationship between relative aggregate stability (standardized MWD) and relative change in 2–5 mm aggregate in the build-up direction.

11
S. Liu et al.
were decomposed (Alvarez et al., 1995, Lorenz and Lal, 2005); more
stable forms of plant C, such as lignin and cellulose, were degraded
during the second phase (16–100 days) (Lorenz and Lal, 2005). Li et al.
(2020) demonstrated that the chemical compositions of wheat and
maize straws were similar at 4 and 6 months, suggesting that a similar
degradation of the various straw components (Baumann et al., 2013)
and microbial community (García-Palacios et al., 2016). Hence, stan­
dardized MWD with milk vetch and maize addition converged in the late
period of incubation. Beyond the chemical composition of residues, soil
microbe varied accordingly with residue decomposition phase. Li et al.
(2020) found an increase in G − bacteria and a decrease in the fungal
proportion in the later period with the degradation of wheat and maize
straws. Therefore, the quality and decomposition phase of the residues
determined the soil aggregation dynamics.
However, the highest standardized MWD was observed in the maize
treatment, followed by milk vetch in Basalt soil. Soil oxides and SOC
acted as essential binding agents in microaggregates and macroaggre­
gates in red soils, respectively (Zhang and Horn, 2001; Peng et al.,
2015b). In this study, the dynamics of the MWD were mainly controlled
by large macroaggregate formation in soils (Fig. 12). Although the soil
sesquioxides varied in the four types of soils (Table 2), these sesqui­
oxides, which mainly facilitated the formation of microaggregates, did
not account for the different change patterns of the MWD in Basalt soil.
Compared with the other three types of soils, the Basalt soil had a higher
pH value (pH = 6.54) because of the lime application in the banana
garden (Table 1). Soil pH is the most important driver of microbial
community composition and diversity (Zhang et al., 2013; Bartram
et al., 2014). Sradnick et al. (2013) indicated that arable soils with
higher pH values resulted in a higher utilisation of amino acid and
carbohydrate-based substrates, which supported microbial growth
(Rousk et al., 2010; Zimmermann et al., 2012). Hence, the abundance of
soil microbes related to maize decomposition is believed to be higher in
Basalt soil in the current study than in the other three types of soils, thus
facilitating a higher standardized MWD with maize application in Basalt
soil (Fig. 3).
Soil aggregation responded more rapidly to microbial activity in
coarse- than fine-textured soils (Fig. 4). Correspondingly, we found that
at a given level of residue incorporation, the relative change of 2–5 mm
in the build-up direction was significantly greater in Sandstone soil
(Fig. 11), thus leading to a faster increase in the standardized MWD in
Sandstone soil relative to the other three types of soils (Fig. S6, sup­
plementary material). In coarse-textured soils, SOC has a greater influ­
ence on structure (Bronick and Lal, 2005). This may be related to the
small specific surface area of soil aggregates in Sandstone soil. Halloysite
predominated in Sandstone soil (Table 1), which provided more sites for
interparticle bonding and clay aggregation (West et al., 2004). However,
a similar result was not observed in Granite soil. Poirier et al. (2014)
reported that unsaturated mineral surfaces in SOC-poor soil favoured
the formation of large macroaggregates and the short-term retention of
C and N in the residues. Compared with Granite soil, Sandstone soil
contained a lower initial SOC content (8.24 vs 3.01 g kg-1), which might
stimulate a sharp increase in the formation of 2–5 mm aggregates with
residue application (Fig. 11). Furthermore, the relative breakdown of
macroaggregates was higher in Granite soil than in the other three types
of soils at a given residue incorporation (Fig. 10), which induced a lower
standardized MWD in Granite soil (Fig. S6, supplementary material).
By taking advantage of the REO labelling method, soil aggregate
formation and breakdown processes are separated. Rather than the
change in the final proportions of macroaggregates, the dynamics of the
MWD were only strongly related to the large macroaggregate formation
rate in this study (P < 0.001, Fig. 12). As indicated in the study by Peng
et al. (2015b), organic matter was a major binding agent for macroag­
gregate formation, but sesquioxides were key agents for microaggregate
formation in red soils. A review paper published by Wiesmeier et al.
(2019) concluded that macroaggregates were formed by binding smaller
aggregates to a central old aggregate with products from residue
12
Geoderma 399 (2021) 115114
decomposition (Six et al., 2002; Blair et al., 2005; Jastrow et al., 2007).
With the incorporation of organic residues, more binding agents facili­
tating aggregate formation were produced (Alami et al., 2000), thus
leading to a remarkable increase in macroaggregate formation relative
to microaggregates (Fig. 11). No other mechanical stresses were applied
under controlled soil moisture conditions during the incubation time,
the relative breakdown of soil aggregates was negligible compared with
the formation processes (Figs. 10 and 11). Therefore, the dynamics of
soil aggregation were controlled by macroaggregate formation more
than breakdown.
5. Conclusions
Residue quality, soil texture and their interactions have critical im­
pacts on soil aggregation. The incorporation of easily decomposable
residue (such as milk vetch in this study) highly increased the formation
but decreased the breakdown of macroaggregates compared with
moderately and slowly decomposable residue (such as maize and
decomposed maize in this study), which was limited by the soil texture
and residue decomposition phase. After residue incorporation, the ag­
gregation in coarse-textured soil had a more rapid response to microbial
activity than in fine-textured soil. Macroaggregate formation dominated
the aggregate build-up and breakdown processes when the soil was
incorporated with organic residues. We conclude that soil aggregation
dynamics are closely related to microbial activity and newly formed
macroaggregates after residue incorporation.
Declaration of Competing Interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influence
the work reported in this paper.
Acknowledgements
This work was granted by the National Natural Science Foundation
of China (41725004, 41571130053). Liu S. was grateful for China
Scholarship Council (Grant No. 201904910375).
Appendix A. Supplementary data
Supplementary data to this article can be found online at https://doi.
org/10.1016/j.geoderma.2021.115114.
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