Applied Soil Ecology xxx (xxxx) xxx–xxx

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Applied Soil Ecology

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

Short communication

Rhizosphere microbial biomass is affected by soil type, organic and water

inputs in a bell pepper system
⁎
Kuan Qina, Xuejun Donga, John Jifonb, Daniel I. Leskovara,
a
Texas A&M AgriLife Research and Extension Center, Texas A&M University, Uvalde, TX 78801, USA
b
Texas A&M AgriLife Research and Extension Center, Texas A&M University, Weslaco, TX 78596, USA

A R T I C LE I N FO A B S T R A C T

Keywords: Effects of environmental and management factors on the diversity and abundance of rhizosphere microbes in
Humic substances unmanaged ecosystems are well established. However, there is limited information about these interactions in
Deficit irrigation vegetable cropping systems. In this two-year field study, we used organic amendments (humic substances, HS)
Clay and sandy soil and water supply (deficit irrigation) as soil management factors to characterize changes in bell pepper rhizo-
Soil microbial
sphere microbial population under two distinct soil types (clay and sandy). Path analysis revealed that soil type
Soil respiration
was the main factor constructing the microbial population. Clay soil indirectly increased bacterial biomass
through the changes in root length, soil pH, P and K contents, while increased fungal biomass was linked with
the enhanced above-ground plant yield. As organic inputs, HS had long-term positive influences on soil microbial
activity through improvements in soil organic carbon content, which was also positively correlated with bac-
terial population. Water input became a less influential factor – deficit irrigation indirectly decreased fungal
biomass but increased soil respiration through the reduction in pepper yield. These findings provide a better
understanding of external factors controlling the abundance of rhizosphere microbiota under intensively man-
aged cropping systems, and potential strategies to improve beneficial microbial diversity and soil health.

1. Introduction
Rhizosphere microorganisms can form symbiotic associations with
host plants, and benefit plant growth and resilience by facilitating re-
source acquisition (e.g. water and nutrients) and pathogen suppression
(Berendsen et al., 2012). The diversity and abundance of rhizosphere
microbial communities, mainly bacteria and fungi, are determined by
plant internal and functional factors such as root growth, respiration
and root-soil chemical exchange (Bulgarelli et al., 2013), as well as
external factors such as soil type, water input, and carbon resource
(Lareen et al., 2016).
The abundance of rhizosphere microbes is energy-driven by the
availability of organic substrates derived from root exudates and
sloughed-off plant cells and tissues. Lange et al. (2015) indicated that
the provision of new and labile carbon substrates had a more important
role than the decomposition of existing soil carbon. Humic substances
(HS), the chemical or biological decomposition products from plant and
animal residuals, are thought to represent a more efficient strategy for
increasing soil organic carbon due to their beneficial effects on plant
root development, shoot growth (or yield) and soil property changes
(Canellas et al., 2015). In recent years, HS have been applied and
evaluated in major agronomic crops (wheat, maize, oat) and high-value
horticultural crops (lettuce, tomato, pepper) (Rose et al., 2014). Being
recognized as promoters for plant root development, HS can increase
the attachment, infection and colonization area for soil microbial
growth. This can improve the synergistic benefits of combined use of
plant growth promoting bacteria (PGPB) and HS (Olivares et al., 2017).
However, compared with the focus on plant growth, less attention has
been given to the changes of rhizosphere microbial population due to
HS application.
It has been noted that soil type and geographical differences are
major determinants influencing the structure of microbial community
(Berg and Smalla, 2009). In a field study where lettuce was grown
under three soil types – sandy, alluvial loam and loess loam, Schreiter
et al. (2014) detected strong effects of soil type on the quantitative
composition of root biomass, as well as rhizosphere bacterial compo-
sition. The role of soil water input on rhizosphere microbial composi-
tion is not fully understood; however, a few studies suggest that water
stress could induce the activity of PGPB and benefit plant resistance to
drought (Rolli et al., 2015). The effects of environmental and man-
agement factors on rhizosphere microbial population in vegetable
cropping systems have received little attention. In order to establish

⁎
Corresponding author.
E-mail address: d-leskovar@tamu.edu (D.I. Leskovar).

https://doi.org/10.1016/j.apsoil.2019.02.024
Received 21 November 2018; Received in revised form 22 February 2019; Accepted 25 February 2019
0929-1393/ © 2019 Published by Elsevier B.V.

Please cite this article as: Kuan Qin, et al., Applied Soil Ecology, https://doi.org/10.1016/j.apsoil.2019.02.024
K. Qin, et al.
Table 1
Basic surface (0–20 cm) soil properties at the study sites in Uvalde and Weslaco, TX.
Soil properties Sand Silt Clay pH EC
(%) (%) (%) (dS/m)
Uvalde 28 25 47 8.2 0.34
Weslaco 63 12 25 7.9 0.48
potential strategies to improve beneficial rhizosphere microbial di-
versity and soil health, a better understanding about how internal and
external factors affect rhizosphere microbial dynamics and composi-
tion, especially under field conditions is needed. Therefore, field ex-
periments were conducted for two years at two Texas locations with
distinct soil types, with or without HS application as organic matter
inputs and subjected to two irrigation levels using an evapotranspira-
tion-based scheduling approach. In order to maintain a long-lasting
effect on rhizosphere properties, a solid lignite-derived HS product with
a relatively slow decomposition rate was used.
2. Materials and methods
2.1. Growth environments, soil types and organic input
Field experiments were conducted in two growing seasons (2016
and 2017) at the Texas A&M AgriLife Research and Extension Centers in
Uvalde, Texas (29.21° N, 99.79° W) represented by a clay soil type, and
Weslaco, Texas (26.16° N, 97.99° W) represented by a sandy soil type
(Table 1). As the weather variables (Fig. A.1) from the two locations
exhibited similar patterns and values, the main difference between the
locations was assumed to be driven by soil type. Lignite-derived solid
humic substances (Novihum Co., Dresden, Germany) were manu-
factured from the reaction with oxygen in aqueous ammonia (ammo-
noxidation procedure), resulting in a product of pH 7.7 and a final
composition of 0.7% fulvic acid, 56.7% humic acid and 24.1% humin,
with 65.8% total C and 3.5% total N. This product was used as organic
inputs by incorporating with surface field soil (0–20 cm) at the rate of
5 t/ha. Due to the extra N contents in HS, adjustments in control were
made using urea fertilizer to ensure that both control and HS-amended
plots received the same amount of N, so that the only difference was the
present of organic matter.
2.2. Plant material and irrigation treatments
Bell pepper (Capsicum annuum cv. Revolution) seedings were ob-
tained from a commercial nursery (Speedling Inc., Alamo, TX) and
transplanted on raised beds with drip irrigation installed at 10–15 cm
below the soil surface in the center bed and covered with plastic mulch.
The experiments were established in a 12 m × 12 m block (total 4
blocks), each block contained 6 rows spaced 2 m apart, in which half (3
rows) of each block was previously amended with HS and the other half
without (control). Using the evapotranspiration (ET) approach (Allen
et al., 1998), bell pepper transplants were subjected to two irrigation
levels in each block: well-watered with 100% ET demands and deficit
(stress) with 50% ET demands. Irrigation was provided based on cal-
culated volumes of crop demands, with the consideration of precipita-
tion and other field conditions such as the percentage of soil covered
with plastic mulch and the flow rate of the drip tape.
2.3. Plant growth responses, soil chemical and biological analysis
In both locations and seasons, harvest started 2 months after
transplanting, and bell pepper yield was recorded. At the end of the
experimental period (4 months after transplanting), part of the plant
roots (0–20 cm depth) were collected using a soil auger (5-cm dia-
meter). Roots were later recovered from the soil cores and scanned
2
Applied Soil Ecology xxx (xxxx) xxx–xxx
NO3-N P K Organic carbon
(mg/kg) (mg/kg) (mg/kg) (%)
18 73 776 1.44
42 29 320 0.19
using an EPSON V700 scanner (Epson, Japan) to measure root length
and surface area using the WinRHIZO software (V5.0, Regent
Instruments, Canada). Thereafter, root dry weight was measured after
oven drying at 75 °C for 2 days.
At the end of the experiment in each season, the same soil auger was
used to collect soil cores within 0–20 cm depth in the plant rhizosphere.
Part of the fresh soil samples collected from the field were immediately
shipped to Earthfort Lab (Corvallis, OR) for microbial biomass analysis
and the rest were used for soil respiration test (Soil CO2-Burst) as de-
scribed by Haney et al. (2008) using the SOLVITA soil respiration box
(Woods End Laboratories, Mt. Vernon, ME). Total bacterial and fungal
biomass were obtained following staining procedures, direct micro-
scope observations, and biomass quantification as described by Qin and
Leskovar (2018). The other part of the soil samples were shipped to the
Soil, Water and Forage Testing Laboratory (Texas A&M AgriLife Ex-
tension Service, College Station, TX) for soil pH, EC, NO3-N, P, K ana-
lysis following the procedures described by Klute (1986). After the two
seasons of growth, soil organic carbon was measured by using the
combustion procedure.
2.4. Statistical analysis
A three-way randomized complete block design with soil type, or-
ganic and water inputs as treatment factors was used in the study. The
data of plant yield, root and soil traits were pooled from the two seasons
and analyzed by ANOVA in R package agricolae (Version 1.2–8); and
multiple comparisons of the means were analyzed by the least sig-
nificant difference (LSD) at α = 0.05. Path analysis was used to illus-
trate the quantitative cause effects of the selected models by using the
lavaan package (Version 0.5–12) in R. The initial path model is de-
scribed in Fig. 1. We hypothesized that these management and en-
vironmental factors would indirectly affect microbial biomass through
the influences on soil chemical properties, plant root and shoot growth.
Management and
environmental factors
Plant Plant Soil
aboveground underground properties
Soil microbial
activity
Fig. 1. Initial path analysis model.
Management factors include organic input (OI) and water input (WI), while soil
type (ST) is considered as environmental factor. Plant aboveground parameter
is mainly represented by yield, while underground is composed of root para-
meters; soil properties include pH, EC and NPK contents; soil microbial activity
contains soil respiration, bacterial and fungal biomass. The covariances among
the soil properties and plant performances are also included but not displayed
in the figure.
K. Qin, et al. Applied Soil Ecology xxx (xxxx) xxx–xxx

Table 2
Bell pepper yield, root parameters, soil chemical properties and rhizosphere microbial activities as affected by management and environmental factors. Management
factors include organic input (OI) and water input (WI), environmental factor includes soil type (ST). Yield is considered as the aboveground trait; underground
parameters include root length (RL), root surface area (RSA), and root dry weight (RDW); soil properties include pH, electrical conductivity (EC), nitrate nitrogen
(NO3-N), available phosphorus (P) and potassium (K). Microbial parameters include soil respiration (SR), bacterial biomass (BB) and fungal biomass (FB).
Source Yield RL RSA RDW pH EC NO3-N P K SR BB FB
(t/ha) (m) (cm2) (g) (dS/m) (mg/kg) (mg/kg) (mg/kg) (mg/kg CO2-C) (μg/g) (μg/g)

OI Control 32.44 10.25 120.72 0.28 8.1 0.49 6.9 b 56 677 22.8 787.7 88.2
HS 34.63 10.45 122.33 0.32 8.1 0.52 9.7 a 57 677 37.5 777.1 111.4
WI 50% 31.40 9.95 115.11 0.28 8.1 0.50 8.7 57 682 29.9 766.3 94.8
100% 35.67 10.75 127.94 0.32 8.1 0.52 7.9 56 671 30.0 798.5 104.8
ST Clay 36.12 a† 11.46 a 138.88 a 0.34 8.1 0.52 8.5 68 a 902 a 42.0 a 1008.6 a 126.9 a
Sandy 30.95 b 9.24 b 104.17 b 0.26 8.0 0.49 8.1 45 b 451 b 18.3 b 556.2 b 72.7 b
P-value OI 0.358 0.812 0.874 0.322 0.355 0.492 0.006 0.601 0.999 0.063 0.854 0.346
WI 0.094 0.345 0.209 0.273 0.800 0.689 0.392 0.601 0.726 0.965 0.579 0.684
ST 0.044 0.011 0.001 0.065 0.056 0.578 0.711 < 0.001 < 0.001 0.003 < 0.001 0.030
OI × WI 0.489 0.234 0.150 0.098 0.933 0.665 0.641 0.741 0.552 0.948 0.140 0.393
OI × ST 0.422 0.050 0.025 0.056 0.275 0.542 0.085 0.164 0.194 0.478 0.172 0.680
WI × ST 0.305 0.199 0.226 0.696 0.555 0.792 0.533 0.492 0.714 0.571 0.822 0.399
OI × WI × ST 0.594 0.515 0.284 0.140 0.933 0.763 0.947 0.510 0.811 0.584 0.956 0.302

†
Different letters within column from same factor indicate significant differences at α = 0.05, according to LSD test.

The observed significant variables were selected based on the ANOVA
to act as indicators of the drivers (endogenous variables). In addition,
direct path from the three factors to soil respiration and microbial
biomass was included to explain the variables not captured by the se-
lected ones – such as soil physical changes. The adequacy of the hy-
pothesized path analysis models was examined by the χ2-test.
3. Results
3.1. ANOVA for bell pepper yield, root and soil parameters
There were significant responses to soil type (ST) and the interac-
tion between organic input (OI) and ST, but there were no significant
interaction effects between the three factors (Table 2). Bell pepper yield
differed significantly between the soil types (P = 0.044), with clay soil
(Uvalde) having 16.7% higher yield than sandy soil (Weslaco). Al-
though not significant, deficit irrigation reduced yield (P = 0.094) by
12.0% compared to the well-watered treatment, while soil OI with HS
application increased pepper yield by 6.8% compared to control. Root
responses also followed a similar pattern as fruit yield, with significant
soil type effects and non-significant OI and soil water input (WI) effects.
Root length and surface area were significantly higher (24.0% and
33.3%, respectively) in clay soil than sandy soil. In addition, there were
significant interactive effects between ST and OI on root parameters. HS
application significantly increased root surface area (RSA, P = 0.025)
and root length (RL, P = 0.050) by 27.1% and 22.6% in sandy soil, but
resulted in slight decreases (14.4% and 12.1%, respectively) in clay soil
compared to control. A strong OI effect on root dry weight (RDW) with
57.7% increase was observed under clay soil, while there was a slight
decrease (10.1%) in sandy soil.
Soil properties collected and measured at the end of the experiments
showed a strong association between soil type and soil pH (P = 0.056),
P and K contents (P < 0.001), with clay soil having higher pH, P and K
concentrations than the sandy soil. OI with HS significantly increased
soil nitrate‑nitrogen (NO3-N) retention compared to control. None of
the factors affected soil electrical conductivity (EC). In terms of rhizo-
sphere microbial population, clay soil exhibited higher soil respiration,
bacterial and fungal biomass than sandy soil. Furthermore, HS appli-
cation increased soil respiration by 64.5% at P = 0.063. Water input
had no effects on soil properties and microbial activity changes.
3.2. Path analysis: from management and environmental factors to
rhizosphere microbial population
Path analysis (Fig. 2) based on data from the two seasons revealed
quantitative relationships from treatment factors to plant, soil proper-
ties and microbial dynamics (including soil respiration, rhizosphere
bacterial and fungal biomass). All the models had good fits (P > 0.9 for
χ2-test) and explained 32% of the variance of soil respiration, 60% of
bacterial and 21% of fungal biomass. Soil respiration was mainly ex-
plained by its positive relationships with organic input and K content,
and negative relationship with yield. Soil K content was significantly
lower in the sandy soil type, while pepper yield was largely associated
with the higher water supply regime (well-watered) and clay soil type.
Rhizosphere bacterial biomass was mostly described by its positive
relationships with soil pH, K content and root growth (root length), and
negative relationships with sandy soil type and P content. Root growth,
soil pH, K and P were all associated with lower sandy content in the
soil. Soil fungal biomass responses were most explained by positive
relationships with yield and negative relationships with soil pH and
sandy content in the soil, although not statistically significant.
4. Discussion
4.1. Soil type was the main environmental factor shaping soil microbial
population
In natural ecosystems, soil type is a less important determinant of
rhizosphere microbial abundance compared to the diversity of plant
species in such native soils. However, in agricultural ecosystems, soil
type becomes a dominant factor, due to frequent soil disturbance and
cultivation of the same crop in different soils (Philippot et al., 2013). In
our study, soil type directly or indirectly shaped the differences in
rhizosphere microbial population through the effects on pepper growth,
soil pH, available P and K contents. Previous studies have shown that
prolific root growth releases more C-rich exudates thus creating re-
source-enhanced micro-niches in the rhizosphere where microbial
community can flourish (Philippot et al., 2009). However, our results
showed that bell pepper root growth had limited effects on rhizosphere
bacterial and fungal biomass. Instead, fruit yield (a key component of
plant shoot biomass) was negatively associated with soil respiration in
clay soil, while fungal biomass was positively associated with higher
yield.
Soil pH could alter the nutrient availability for microbiota in the
rhizosphere (Philippot et al., 2013). Fierer and Jackson (2006) found
that bacterial diversity was highest in neutral soils and lower in acidic
soils. The soil types in this study were slightly alkaline, with the clay
soil having a higher pH and supporting higher bacterial abundance but
decreased fungal population compared to the sandy soil. The soil type
effects on microbial diversity could also reflect outcomes of natural

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K. Qin, et al. Applied Soil Ecology xxx (xxxx) xxx–xxx

Fig. 2. Path analysis model explaining the manage-

WI ST OI a ment and environmental factors affecting soil re-
spiration (a, χ192 = 9.57, P = 0.963), rhizosphere
bacterial (b, χ202 = 9.46, P = 0.977) and fungal
-0.849** biomass (c, χ202 = 9.66, P = 0.974). External fac-
0.213* -0.285* -0.274* -0.781** 0.284** tors were treated as binary variables – for the or-
-0.187
ganic input (OI), 0 represents control and 1 re-
presents HS application; for the water input (WI), 0
RL Y pH P K N means deficit irrigation (50% ET) and 1 means well-
watered (100% ET) treatment; for the soil type (ST),
0.251*
-0.097 0 denotes clay soil and 1 denotes sandy soil.
-0.070 -0.086 Numbers next to the arrows indicate standardized
path coefficients and the significance is marked by
-0.291* 0.551* * < 0.05 or ** < 0.01. The numbers next to the
-0.081 -0.009 endogenous microbial variables represent their ex-
plained variance (R2). RL: root length; Y: yield; pH,
Soil respiration 0.32
N, P, K: soil properties from soil samples. All the
models contain significant (< 0.05) covariances
between Y and RL, Y and pH, Y and P, Y and N, N
and pH, N and K, P and pH, P and K. All error terms
WI ST OI b are not displayed in order to improve the readability
of the figure.

-0.849**
0.213* -0.285* -0.274* -0.781** 0.284**
-0.187

RL Y pH P K N

0.231* -0.789** -0.473**

0.112 -0.056 0.243 -0.081

Soil bacterial biomass 0.60

WI ST OI c

0.213* -0.285* -0.274* -0.849**

-0.781** 0.284**
-0.187

RL Y pH P K N

-0.254 -0.397 -0.040

-0.003 0.162 -0.097 -0.042

Soil fungal biomass 0.21

competitive interactions between bacteria and fungi. The bacteria in
this study were more abundant than fungi, which resulted in a lower F/
B (fungi to bacteria) ratio with 0.13 in both soil types. Soil type also
significantly affected soil P and K retention, and clearly clay soil had
more nutrient contents than sandy soil (Table 2). These two nutrients
have previously been shown to be similarly partitioned in explaining
the soil microbial diversity (Su et al., 2015). In our study, P retention
had a negative influence on rhizosphere bacteria, while K was posi-
tively associated with soil respiration and bacterial population. Nu-
merous phosphate- and potassium- solubilizing microorganisms that
help plants mobilize insoluble P and K are affected by soil available P
and K contents (Adhya et al., 2015; Bahadur et al., 2016). Our results
suggest that there might be competition between these functional mi-
crobes.
In addition to impacting soil microbial composition indirectly
through the changes in plant root growth, yield and basic soil proper-
ties, soil type also influenced rhizosphere bacterial and fungal popula-
tion via other factors (Fig. 2). These could include soil porosity, ag-
gregation, hydraulic conductivity and other physical traits. This study
recognized that more parameters had to be investigated and added in
the future in order to clearly understand why soil type became the main
factor in differentiating soil microbiota.

4
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4.2. Humic substances as organic inputs may have a long-term influence on being more sensitive to water input changes as compared to fungi. From
soil microbial activity the path analysis, we found that higher water input contributed to the
increase of pepper yield, but negatively affected soil respiration, which
The presence of HS could be used as carbon substrate to attract and possibly due to the reduced bacterial biomass and other microbial ac-
support the bacterial population growth in the rhizosphere (Olivares tivity except for fungi. Instead, extra soil moisture positively induced
et al., 2017). However, our study did not show a significant impact of fungi abundance through improvement in plant aboveground perfor-
HS input on root growth or rhizosphere microbial biomass, except for mance, which is also in agreement with Ma et al. (2016). Overall, water
the enhancement of soil respiration. Soil respiration reflects the CO2 input was the least influential management factor that shaped the rhi-
efflux generated by decomposition of rhizosphere organic matter from zosphere microbial biomass. A slight water stress may achieve a more
soil biota, which has been used as an indicator to quantify the impacts abundant microbial community compared to non-limited irrigation.
of various environmental and management factors on microbial activity
and soil health (Haney et al., 2008). In the current study, HS application
5. Conclusions
seemed to have beneficial influences on microbial activity, but this was
not reflected by the selected plant growth or soil related variables in the
In this two-year field study conducted in a bell pepper production
path analysis. At the end of the experiments, HS treated soil retained
system, we examined the strength of two management factors (organic
more N in the form of nitrate than control plots. This enhanced reten-
and water inputs) and one environmental factor (soil type) in influen-
tion capacity is important to prevent leaching losses and thus can serve
cing rhizosphere microbial population. Soil type was the main factor
as nutrient storage for succeeding crops. Reported effects of N supply on
shaping the microbial population, with clay soil having more microbial
soil microbial biomass are mixed. For instance, Li et al. (2015) showed
activity than sandy soil. Differences in soil nutrient also reinforced the
that excessive N supply had no effects on microbial biomass, whereas
significant role of soil type in explaining the microbial activity changes.
other study indicated that extra N decreased fungal activity (Allison
Organic input represented by HS application significantly influenced
et al., 2007).
soil nitrate‑nitrogen (NO3-N) retention, as well as soil respiration. Over
Soil organic carbon (SOC) had higher positive correlation with
time, the soil organic carbon incorporated by HS induced positive
bacteria (P < 0.001) than with fungi (P = 0.046), but uncorrelated
correlations with bacterial and fungal biomass, which could benefit
with soil respiration (Fig. A.2). Although clay soil had higher SOC than
sustainable agriculture – by decreasing exogenous mineral nutrients
sandy soil, the SOC was significantly higher in HS treated soil compared
supply and increasing endogenous organic matter inputs, soil biological
to control regardless of soil type (HS 1.65% vs. control 1.37% in clay
activity becomes more vigorous in the rhizosphere. The effects of soil
soil and 0.64% vs. 0.43% in sandy soil). This suggested the positive
water input on rhizosphere microbial biomass were not remarkable, but
interactions between HS and heterotrophic or autotrophic bacteria,
could significantly affect plant yield. While high irrigation may inhibit
which can benefit from the consumption of certain organic compounds
soil respiration, moderate water stress achieved by deficit irrigation can
such as organic acids or amino acids associated with increased organic
up-regulate the rhizosphere microbial activities and conserve water
carbon supply due to HS application (Valdrighi et al., 1996). However,
resources. Discovery of key factors that affect the rhizosphere microbial
research on specific types of microbes and their influences to the soil
population is essential to guide management practices for improving
environment and rhizosphere dynamics is still lacking and needs fur-
mutually favorable relationships between microbes and plants, as well
ther investigation.
as soil ecosystem functions.
4.3. Water input affected soil respiration
Acknowledgements
Water input can affect soil oxygen pressure (Centenaro et al., 2017)
and subsequently, root water uptake and respiration. Root growth and This work was supported by the Novihum Technologies GmbH,
activity can in turn impact microbial activity and respiration through Germany, and partially by the Texas A&M AgriLife Cropping System
provision of carbon exudates (Philippot et al., 2013). Based on a meta- program. We thank Dr. Yahia Othman, Dr. Desire Djidonou, Manuel
analysis (Manzoni et al., 2012), soil microbial activity in the commu- Figueroa Pagan and Maria Andrea Macias for their assistance in this
nity-level decreased with the reduction of soil moisture, with bacteria study.

5
K. Qin, et al. Applied Soil Ecology xxx (xxxx) xxx–xxx

Appendix A

Max Mean Min

40
Temperature (°C)

30

20

10

0
100
Relative humidity (%)

75

50

25

0
Daily light integral (mol ⋅ m−2 ⋅ day−1)

100

75

50

25

15
Precipitation (mm)

10

0
Aug Oct Dec Feb Apr Jun Aug Oct Dec Feb Apr Jun
2016 2017 2016 2017

Fig. A.1. Weather data from Uvalde and Weslaco, TX in 2016 and 2017.
Letter “S” indicates the start time of the experiments (after transplanting), letter “E” indicates the end time of the experiments (after final harvest). During the
experimental period, the seasonal mean temperature, relative humidity, daily light integral and total precipitation were 23 °C, 73%, 66 mol·m−2·day−1 and
426 mm, respectively in 2016 Uvalde; 26 °C, 67%, 86 mol·m−2·day−1 and 210 mm, respectively in 2017 Uvalde; 26 °C, 72%, 70 mol·m−2·day−1 and 155 mm,
respectively in 2016 Weslaco; nd 30 °C, 72%, 88 mol·m−2·day−1 and 126 mm, respectively in 2017 Weslaco.

6
K. Qin, et al.
Fig. A.2. Linear regression between soil carbon content and soil respiration, rhizosphere bacterial and fungal biomass in soil samples collected at the end of the
second-year experiment.
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