Biochar

https://doi.org/10.1007/s42773-019-00009-2

REVIEW

Response of microbial communities to biochar‑amended soils:

a critical review
Kumuduni Niroshika Palansooriya1 · James Tsz Fung Wong2 · Yohey Hashimoto3 · Longbin Huang4 ·
Jörg Rinklebe5,6 · Scott X. Chang7 · Nanthi Bolan8 · Hailong Wang9,10 · Yong Sik Ok1 

Received: 18 March 2019 / Accepted: 24 March 2019

© Shenyang Agricultural University 2019

Abstract
Application of biochar to soils changes soil physicochemical properties and stimulates the activities of soil microorganisms
that influence soil quality and plant performance. Studying the response of soil microbial communities to biochar amendments
is important for better understanding interactions of biochar with soil, as well as plants. However, the effect of biochar on soil
microorganisms has received less attention than its influences on soil physicochemical properties. In this review, the following
key questions are discussed: (i) how does biochar affect soil microbial activities, in particular soil carbon (C) mineralization,
nutrient cycling, and enzyme activities? (ii) how do microorganisms respond to biochar amendment in contaminated soils?
and (iii) what is the role of biochar as a growth promoter for soil microorganisms? Many studies have demonstrated that
biochar-soil application enhances the soil microbial biomass with substantial changes in microbial community composition.
Biochar amendment changes microbial habitats, directly or indirectly affects microbial metabolic activities, and modifies the
soil microbial community in terms of their diversity and abundance. However, chemical properties of biochar, (especially
pH and nutrient content), and physical properties such as pore size, pore volume, and specific surface area play significant
roles in determining the efficacy of biochar on microbial performance as biochar provides suitable habitats for microorgan-
isms. The mode of action of biochar leading to stimulation of microbial activities is complex and is influenced by the nature
of biochar as well as soil conditions.

Keywords  Carbon cycling · Microbial habitat · Microorganisms · Nutrient cycling · Enzyme activity

* Yong Sik Ok of Foundation Engineering and Water and‑Waste

yongsikok@korea.ac.kr
1
Korea Biochar Research Center, O‑Jeong Eco‑Resilience
Institute (OJERI), Division of Environmental Science
and Ecological Engineering, Korea University, Seoul 02841,
Republic of Korea
2 Edmonton, AB T6G 2E3, Canada
Department of Civil and Environmental Engineering,
The Hong Kong University of Science and Technology,
Hong Kong SAR, China
3 9
Department of Bioapplications and Systems Engineering,
Tokyo University of Agriculture and Technology, Tokyo,
Japan
4
Environment Centres, Sustainable Minerals Institute, The
University of Queensland, Brisbane, QLD 4072, Australia
5
Laboratory of Soil‑ and Groundwater‑Management,
School of Architecture and Civil Engineering, Institute
Management, University of Wuppertal, Pauluskirchstraße 7,
42285 Wuppertal, Germany
6
Department of Environment, Energy and Geoinformatics,
Sejong University, Seoul 05006, Republic of Korea
7
Department of Renewable Resources, University of Alberta,
8
Global Centre for Environmental Remediation, University
of Newcastle, Callaghan, NSW 2308, Australia
School of Environmental and Chemical Engineering, Foshan
University, Foshan 528000, Guangdong, China
10
Key Laboratory of Soil Contamination Bioremediation
of Zhejiang Province, Zhejiang A&F University,
Hangzhou 311300, Zhejiang, China

13
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 Biochar

1 Introduction vital role in soil ecosystem functions and services (for

Biochar exhibits a range of physicochemical properties
depending on the type of feedstock, pyrolysis conditions,
and modification techniques (such as activation, magnetic
modification, and acid/base treatments) (Lou et al. 2016;
Mohan et al. 2014; Rajapaksha et al. 2016). Even though
the chemical and physical properties of biochar have
been well studied, the biochar’s effects on the biologi-
cal functions of soil is poorly understood. Understanding
the impacts of biochar on soil biological functions would
require long-term monitoring and investigation of changes
in biological properties in biochar-amended soils. Biochar
application would have complex effects on soil biological
properties, including direct/indirect interactions among
abiotic and biotic factors and processes in the soil (Zhu
et al. 2017). Over the past couple of decades, studies have
revealed that biochar-soil application could improve soil
biological properties by enhancing soil microbial func-
tional activities and altering community structure (Li et al.
2018), (Fig. 1). Moreover, biochar effects on soil biolog-
ical properties as affected by other soil organisms and
plants were also studied (Palansooriya et al. 2019; Paz-
Ferreiro et al. 2014). Since soil microorganisms play a
example, driving biogeochemical cycles, suppressing path-
ogens, and maintaining soil quality and health), the next
milestone of biochar research should focus on understand-
ing the long-term impact of biochar application on soil
biota together with soil health. It is necessary to explore
the capacity of biochar to improve soil quality under future
environmental changes (Griffiths and Philippot 2013).
The soil is home to microorganisms such as bacteria,
fungi, nematodes, algae, archaea, actinomycetes, bacte-
riophages, and protozoa (Paul 2014). These microbes are
involved in many beneficial soil functions, such as nutrient
recycling, organic matter decomposition, soil-structure for-
mation, secretion of plant growth promoters, degradation of
organic contaminants, and suppression of pests and diseases
(Kirchman 2018). Soil microbial functional activities and
community structure can be considered as useful indica-
tors in assessing the impacts of biochar on soil biological
properties. Some parameters for example, basal respiration,
nitrogen (N) mineralization, N ­ 2 fixation, and enzyme (e.g.,
dehydrogenase) activities are used to determine microbial
functional activities in the soil (Paz‐Ferreiro and Fu 2016),
whereas phospholipid fatty acid (PLFA), denaturing gradi-
ent gel electrophoresis (DGGE), temperature gradient gel
electrophoresis (TGGE), polymerase chain reaction (PCR),

ATTRIBUTES RESPONSE OUTCOMES

Supply of nutrients; Enhanced soil microbial community

+ 2+ 2+ 3- +
K , Mg , Ca , PO4 , NH4 abundance, acvity, and diversity Facilitate biochemical cycling

Enhanced water holding Improve soil structure

capacity through microaggregates

Changes in soil pH,

electrical conducvity Improve soil enzymes

Biochar-soil Contaminant degradaon

Biochar applicaon
Transform into less toxic
forms by oxidaon,
reducon, and hydrolysis

High surface area and porous

structure provide beer Contaminant immobilizaon
habitats
-Sorpon into cells
Low temperature-biochar -Formaon of complexes/precipitates
provides readily available C as
an energy source Promoon of plant

Release of microbial Control of potenal

inhibitors pathogens

Fig. 1  Schematic diagram showing the effect of biochar on soil microorganisms and microbial responses following biochar application

13
Biochar
and DNA and RNA analyses are some of the useful methods
for determining microbial community structure (Nannipieri
et al. 2003).
After the review by Lehmann et al. (2011), many stud-
ies have reported biochar-induced changes in soil micro-
bial properties, including microbial biomass, community
composition and diversity, specific functional groups, and
biogeochemical processes (Awad et al. 2018; Li et al. 2018;
Xu et al. 2018b). However, it is difficult to draw conclu-
sions about the mechanisms of biochars regulating micro-
bial structure and functions in the soil (Table 1). This is
not unexpected due to the wide range of biochar properties,
soil types, and experimental conditions that affect micro-
bial properties. The physicochemical properties of biochar
(Table 2) greatly impact soil biological properties following
its application. For example, the increase in soil pH, dis-
solved organic C content, carbon-to-nitrogen ratio (C:N),
and ­K+ concentrations in biochar-amended soils was found
to alter microbial community structure in different soils
(Choppala et al. 2016; Muhammad et al. 2014; Quilliam
et al. 2013; Wong et al. 2019). Moreover, Farrell et al. (2013)
reported that the application of biochar improved the soil
labile C content and soil pH and altered soil microbial com-
munity structure. Nevertheless, the highly porous structure
of biochar provides habitats for microorganisms to colonize,
permitting their growth in the soil environment (Awad et al.
2018; Lehmann et al. 2011; Quilliam et al. 2013; Ye et al.
2017). However, neutral/negative impacts of biochar on
soil biological properties have also been documented. For
example, Elzobair et al. (2016) found that the application
of biochar to an Aridisol had no effect on soil extracellular
enzyme activity, microbial community structure, microbial
biomass, and nor arbuscular mycorrhizal fungi root coloni-
zation. The authors suggested that biochar’s effect on micro-
bial communities largely depended on biochar application
rates, biochar and soil types. The changes in microbial func-
tions and community structure could only occur when the
application rates of biochar are high enough to significantly
alter soil moisture retention capacity, pH conditions, and/
or nutrient concentrations, which are the most critical fac-
tors influencing microbial diversity and composition in the
soil (Gomez et al. 2014). For instance, increasing biochar
application rate from 0.5 to 5.0% (w/w) reduced the total
PLFA content and the relative abundance of bacteria in the
soil (Wang et al. 2015). Furthermore, it was suggested that
smaller amounts of biochar application (0.5% (w/w)) could
increase the activities of soil extracellular enzymes such as
ß-D-cellobiosidase, ß-glucosidase, and N-acetyl-ß-glucosa-
minidase (Wang et al. 2015).
Soil condition plays a key role in determining microbial
community composition in biochar-amended soils. For
instance, Awad et al. (2018) observed lower PLFA bio-
mass in biochar-treated soils under oxidized conditions,
whereas PLFA biomass was increased when the soil was
under reduced conditions. Microbial communities may not
be stimulated if biochar could not provide sufficient labile
C or N substrates, and if the biochar application rate is not
sufficient to affect the soil pH.
The recent increasing trend in the number of studies on
biochar effects on soil biota (Fig. 2) justifies a comprehen-
sive review on this topic and proposes much needed future
research questions and directions. Despite the increasing
research on biochar–microbial relationships, mechanistic
studies have remained few, focusing on discerning specific
roles of biochar in microbial processes in biochar-amended
soils. The previous reviews have illustrated key aspects of
biochar from production methods, physical and chemical
characterization, to interactions with soil physicochemical
properties (Jeffery et al. 2011; Lehmann et al. 2006b; Sohi
et al. 2010). Biochar had been extensively applied as amend-
ments for environmental remediation, such as removal and
degradation of inorganic and organic contaminants from soil
and water (Ahmad et al. 2014; Mohan et al. 2014).
Nevertheless, the knowledge on the influences of biochar
on soil biota has remained limited and poorly integrated.
Thus, the objectives of this review are to discuss the (i) role
of biochar as a growth promoter for soil microorganisms, (ii)
impacts of biochar on the soil microbial activities, and (iii)
responses of microbial communities to biochar amendment
in contaminated soils. This review will enable us to identify
research gaps in this area.
2 Biochar as a growth promoter for soil
biota
Modification of soil physicochemical conditions may
positively or negatively affect soil microbial community
structure and functions, through altering their habitats.
These changes in soil properties are the direct and primary
growth promoter for soil microbes. Improvements in soil
properties, such as increase in soil pH, soil–water retention
capacity, and nutrient [e.g., C, N, phosphorus (P), potas-
sium (K), magnesium (Mg), and calcium (Ca)] availability
(Li et al. 2019) could promote soil microbial growth. The
available nutrient composition and concentration in biochar
are governed by the feedstock and production conditions,
such as production temperature and retention time. In gen-
eral, increase in production temperature increases biochar
pH, total surface area, carbon-to-oxygen ratio (C:O), and
C:N, while reducing dissolved organic matter and C contents
(Crombie et al. 2013, Huff et al. 2014, Uchimiya et al. 2013).
In addition, biochar also supports microbial communities by
providing labile C substrates for degradation (Smith et al.
2010). In this section, we summarize the impact of physico-
chemical properties of biochar on soil biota. It shows some
13
 Table 1  Effects of biochar application on soil microbial community/structure
Biochar feedstock Production Dose (%) Soil type Soil microbial response(s) Possible mechanism(s) involved References
temperature

13
(°C)

Willow leaves 470 0, 0.5, 2 Flinty clay loam Increased total bacteria BC application affects net N mineralization Prayogo et al. (2014)
by 28%, Gram-negative processes
bacteria by 27% and act-
inobacteria by 62%
Maize straw 400 0, 0.7, 2.2, 3.7 Silt loam BC application affected SOC and ­KMnO4-oxidizable C (­ KMnO4–C) Luo et al. (2017)
AMF and the ratio of determine the soil microbial community
AMF/saprotrophic fungi composition
Oak and hickory hardwood 500 0.17 Aridisol BC did not affect the micro- Little biochar C was available for microbial Elzobair et al. (2016)
sawdust bial community structure, degradation
biomass, extracellular BC did not improve total N, N­ H4-N, ­NO3-N,
enzyme activities, and or available P in soil
AMF root colonization
Rice straw 550 0.365 Clay loam BC had a slight influence BC enhanced further mineralization of Pan et al. (2016)
on microbial biomass SOC and high C availability stimulated
Relative abundance of microbial growth
microbial community had
a positive correlation with
C availability. Increased
population of Gram-nega-
tive bacteria
Bull manure + dairy 500 0–10 Fertile Mollisol Increased microbial biomass NA Kolb et al. (2009)
manure + Pinus spp. and microbial activity
Swine manure and willow 350 0–10 Temperate sandy loam soil Increased microbial biomass NA Ameloot et al. (2013)
wood C and dehydrogenase
activity
Increased abundance of
Gram-positive and Gram-
negative bacteria
Pinus radiata NA 10 Silt-loam soil Promoted phosphate solubi- NA Anderson et al. (2011)
lizing bacteria, increased
the abundance of bacterial
families that can degrade
more recalcitrant C com-
pounds, decreased bacte-
rial plant pathogens
Wheat straw 350–550 0, 1.5, 3 Slightly acid rice soil Increased bacterial gene BC addition changed soil properties, such Chen et al. (2013)
abundance and decreased as enhanced SOC and pH. It also changed
fungal gene abundance soil biochemical processes and, therefore,
Significant increase in the the soil enzyme activities
activity of dehydrogenase
and alkaline phosphatases
Biochar
 Table 1  (continued)
Biochar

Biochar feedstock Production Dose (%) Soil type Soil microbial response(s) Possible mechanism(s) involved References
temperature
(°C)

Corn straws 500 0.5 Sandy loam soil Increased microbial biomass Inhibitory effect on SOC decomposition Lu et al. (2014)
especially Gram-positive
bacteria
Promoted fungi growth
Yeast-derived BC NA 30 Arable and forest soil Promoted fungi in the soil Enhanced nutrient levels in soil Steinbeiss et al. (2009)
Glucose-derived BC NA 30 Arable and forest soil Gram-negative bacteria uti- NA Steinbeiss et al. (2009)
lized the glucose-derived
BC
Maize straw 450 0–5.0 Fluvo-aquic soil, light loamy Increased activity of soil Significantly increased SOC, total N, and Wang et al. (2015)
soil extracellular enzymes exchangeable K
which involved in C, N,
and S cycling
Sawdust 550 5.2 Sandy loam Reduced total PLFA Low available C due to high sorption affin- Awad et al. (2018)
biomass at high redox ity of BC for SOM and DOC
potential Imitation of N content in treated soils
Increased total PLFA Increased OM content and available C,
biomass at low redox increased soil pH
potential
Oak pellet 550 1–20 Colorado, USA; Sandy loam 40% decrease in total Larger quantities of BC modified the soil Gomez et al. (2014)
PLFA-C structural properties
Iowa, USA; Sandy loam 64% decrease in total BC could add some toxic compounds which
PLFA-C cause negative effects on microorganism
Michigan, USA; Clay 56% decrease in total
PLFA-C
Minnesota, USA; Clay loam 61% decrease in total
PLFA-C
Compost NA 2 Mixture of compost and In DGGE profile, a number BC provided better space and good elec- Chen et al. (2017b)
river sediment of bands were significantly tronic transmission which were beneficial
increased in BC amended for microorganisms to use of the refrac-
treatment tory substances in treated sample
Raw rice husk 500 2 River sediment 16S rRNA sequencing anal- Addition of BC changes the Liu et al. (2018)
ysis showed that the appli- physicochemical properties of the sediment,
cation of BC adversely which has a negative effect on the sedi-
affected the behavior of ment microorganisms
indigenous microbes,
reduced abundance, and
diversity of bacteria

AMF Arbuscular mycorrhizal fungi, BC biochar, HM heavy metal, NA not available, OM organic matter; PAH polyaromatic hydrocarbons, SOC soil organic carbon

13
 Table 2  Selected physicochemical properties of biochar produced under different temperatures with different feedstock types [data obtained from UC Davis Biochar Database (2019)]
Feedstock Pyro. Temp. Total ash content Volatile matter C N H O PH EC Cation exchange Total surface area
capacity

13
°C % dS ­m−1 cmolc kg−1 m2 g−1

Peanut hull 400 8.20 38.40 74.80 2.70 4.50 9.70 7.90 0.54
Peanut hull 500 9.30 18.10 81.80 2.70 2.90 3.30 8.60 1.22
Poultry litter 350 35.90 36.70 46.10 4.90 3.70 8.60 8.70 1.10
Poultry litter 700 52.40 14.10 44.00 2.80 0.30 0.01 10.30 9.00
Soft wood 700 6.40 58.10 0.41 4.16 31.66 6.80 66.96 2.03
Turkey litter 700 64.00 15.60 0.78 0.83 4.37 10.90 24.45 21.77
Walnut shell 900 40.40 55.30 0.47 0.89 1.55 9.70 33.40 227.10
Hog waste 650 2.40 68.20 0.51 3.66 26.82 7.50 26.21 25.15
Wood feedstock 510 3.00 83.90 0.36 1.88 19.82 7.30 12.00 165.80
Wood feedstock 410 2.60 65.70 0.21 4.38 23.52 7.10 10.00 2.82
New earth pine 600 17.00 71.20 0.91 2.88 11.61 7.90 3.18 4.97
Cottonwood 4.20 82.50 0.49 1.64 5.61 9.50 16.47 300.00
Corn stover 700 58.80 7.60 33.50 1.00 5.10 29.00
Corn stover 450 58.00 12.70 33.20 1.40 8.60 12.00
Bubinga wood 250 0.90 66.40 34.90 5.40
Bubinga wood 400 1.80 41.10 17.60 6.10
Bubinga wood 525 1.20 35.00 11.40 500.90
Bubinga wood 650 1.30 22.30 13.80 548.90
Laurel oak wood 250 1.40 66.00 41.50 1.80
Laurel oak wood 400 2.60 51.90 25.90 2.20
Laurel oak wood 525 6.80 36.40 21.70 38.20
Laurel oak wood 650 3.70 20.70 18.20 218.70
Loblolly pine wood 250 0.30 61.10 38.20 139.70
Loblolly pine wood 400 0.50 58.60 25.90 2.90
Loblolly pine wood 525 1.20 25.70 14.00 206.10
Loblolly pine wood 650 1.10 25.20 12.20 393.90
Sugarcane trash 500 68.00 1.20 9.60 4.80 40.00
Sugarcane bagasse 500 65.00 1.10 8.40 0.18 3.50
Rice straw 200 11.50 46.70 0.79 5.00 34.30 2.10
Rice straw 350 31.00 46.90 2.08 1.34 12.30 20.60
Rice straw 500 48.40 40.70 1.18 0.15 4.14 128.00
Wheat straw 350 55.00 1.10 5.90 65.00
Wheat straw 550 69.00 1.10 8.50 205.00
Pine needle 350 67.00 2.80 6.70 30.00
Biochar
 Table 2  (continued)
Biochar

Feedstock Pyro. Temp. Total ash content Volatile matter C N H O PH EC Cation exchange Total surface area
capacity
°C % dS ­m−1 cmolc kg−1 m2 g−1

Pine needles 550 76.00 2.50 4.60 83.00

Wheat straw 525 19.80 69.60 1.50 2.10 7.10 10.10 0.60
Wheat straw 525 21.60 49.30 1.20 3.70 24.10 6.80 1.60
Wheat straw 550 26.40 49.60 1.10 2.90 20.00 7.10 2.40
Sawdust wood 500 75.80 0.30 8.03 8.42 203.00
Grass 500 62.10 2.83 1.65 17.80 3.33
Chlorella algae 500 39.30 3.37 5.35 12.60 2.78
Eucalyptus wood 600 1.74 2.19 81.03 1.07 10.40 334.56
Grass straw 200 5.70 70.70 47.20 0.61 7.11 45.10 3.30
Grass straw 300 9.40 54.40 59.70 1.02 6.64 32.70 4.50
Grass straw 400 16.30 26.80 77.30 1.24 4.70 16.70 8.70
Grass straw 500 15.40 20.30 82.20 1.09 3.32 13.40 50.00
Grass straw 600 18.90 13.50 89.00 0.99 2.47 7.60 75.00
Grass straw 700 19.30 9.10 94.20 0.70 1.53 3.60 139.00
Palm wood 300 59.22 1.33 6.47 1.33 3.10
Palm wood 600 64.32 0.67 10.13 2.36 196.40
Pine wood 300 71.90 0.25 4.63 0.07 9.60
Pine wood 600 72.35 0.21 6.75 0.06 112.90
Nutshell 300 60.60 0.35 6.88 0.72 0.04
Nutshell 600 71.27 0.66 9.85 1.60 59.50
Nutshell 300 69.63 1.79 6.11 0.31 1.70
Nutshell 600 79.77 0.62 9.06 1.01 261.60

13
 Biochar
Fig. 2  Publications on “bio-
char” and “microbial” from
1990 to 2018 (SCOPUS data
base)
Number of publications (biochar+microbial )
selected physicochemical properties of biochar produced
under different pyrolysis temperature with different feed-
stock types are summarized in Table 2.
The highly porous nature of biochar and its high surface
area are favorable habitats for soil biota (Jaafar et al. 2014,
2015; Ye et al. 2017). The specific area and the porosity of
biochar strongly depend on the type of feedstock as well
as the production temperature. High temperature promotes
organic matter and water volatilization during the thermal
decomposition process and results in large pores (Brewer
and Brown 2012). Figure 3 shows the highly porous struc-
ture of biochar through scanning electron microscopy
(SEM), in which the feedstocks are mostly plant origin, and
the biochars were produced at 600 °C and 850 °C. The bio-
char has macropores (> 200 nm), mesopores (2–50 nm), and
micropores (< 2 nm), according to the IUPAC classification
(Lehmann and Joseph 2015; Zhao et al. 2013). These pores
serve as habitats for diverse soil biota (Jaafar et al. 2014;
Pietikäinen et al. 2000; Quilliam et al. 2013; Warnock et al.
2007), such as bacteria (size within the rage of 0.3–3 mm),
ectomycorrhizal fungi, ericoid mycorrhizal fungi, and arbus-
cular mycorrhizal fungi (AMF) (2–80 mm) (Warnock et al.
2007). Bacteria and the hyphae (< 16 µm) of different fungi
colonize the biochar pores, gaining physical protection from
predators (such as mites and nematodes) in the soil (Ezawa
et al. 2002; Pietikäinen et al. 2000; Saito 1990). Coloni-
zation of biochar pores by AMF has also been reported
(Ezawa et al. 2002; Gaur and Adholeya 2000; Saito 1990).
In addition, water molecules and soluble substances (e.g.,
acids, alcohols, aldehydes, ketones, and sugars) involved in
microbial metabolism would be stored in mesopores and
micropores (Brewer and Brown 2012; Lehmann and Joseph
13
140
[biochar+ microbial (Soil fertility/Soil biota /Soil respiration/Contaminated
250 Number of publication
soil/Soil enzyme activity/Fung+ bacteria/Plant growth promoter)]
Soil fertility
Soil biota 120
Soil respiration
200 Contaminated soil
Soil enzyme activity 100
Fung+bacteria
Plant growth promoter
Number of publications
150 80
60
100
40
50
20
0 0
2007 2008 2009 2010 2011 2012 2013 2014 2015 2016 2017 2018
Year
2015). As a result, biochar may regulate the abundance and
activities of certain microbes which could take advantage of
the physical properties of biochars.
Surface area is another important biochar property that
influences the growth of soil microbes (Jaafar et al. 2015).
It is generally higher than that of the corresponding feed-
stock materials and also the soil. When compared with slow
pyrolysis, fast pyrolysis can increase the surface area of
biochar (Chintala et al. 2014a; Mohanty et al. 2013). The
surface areas of soybean stover biochar produced at 300 and
700 °C were 5.61 and 420.33 m2 g−1, while pine needles
biochars produced at the same temperatures showed 4.09
and 390.52 m2 g−1 of surface areas, respectively (Ahmad
et al. 2016). Thus, depending on the pyrolyzing temperature
and the type of feedstock, increased porosity and surface
area provide a greater access for microbial colonization.
Increased specific surface area and porosity of biochar are
related to increased water holding capacity across different
soil types, providing improved water retention for microbial
activities and growth (Ahmad et al. 2014; Mohamed et al.
2016; Wong et al. 2017). An incubation study conducted
with woody biochar revealed better hyphal colonization on
the biochar surface via SEM (Jaafar et al. 2015). In addi-
tion, the study suggested that the enhanced surface area and
the high porosity of biochar provided potential habitats for
soil microorganisms (Jaafar et al. 2015), (Figure 4 shows
the SEM images of fungal hyphae grown on peanut shell
biochar surface).
Soil chemical properties, particularly soil pH is one of the
governing factors of soil microbial abundance and activity,
which may be significantly modified by biochars if a suffi-
cient amount of biochar is added to the soil (Lehmann et al.
Biochar
Fig. 3  Scanning electron microscopy (SEM) images of various types
of biochars (a The mesquite wood chip biochar produced at 850 °C;
b 40% food waste + 60% wood biochar produced at 850 °C; c Wood
2011). The presence of negatively charged hydroxyl, car-
boxyl, and phenolic groups on the surface of biochar makes
them alkaline (Brewer and Brown 2012; Chintala et  al.
2014b). In addition, carbonates, bicarbonates, and silicates
in biochar can bind with the hydrogen ion in soil–water. It
­ + concentration and, therefore, an increase in
reduces the H
pH value (Gul et al. 2015). The alkalinity of biochar is also
positively correlated with the production temperature (Gul
et al. 2015) and related to the feedstock type. Therefore,
biochar application usually increases soil pH (Masto et al.
2013; Stewart et al. 2013; Wong et al. 2017; Xu et al. 2014).
The addition of corn straw-derived biochar increased the
soil pH and electrical conductivity (EC) and decreased N
leaching; as a result, changes were observed in the commu-
nity composition of bacteria (Xu et al. 2016). In addition,
the biochar application rate was found to be positively cor-
related with bacterial diversity in the biochar-amended soil
(Xu et al. 2016). However, Luo et al. (2017) reported that
the addition of maize straw biochar did not increase the soil
chips biochar produced at 850  °C, and d dairy manure biochar pro-
duced at 600  °C) (images provided by Korea Biochar Research
Center at Korea University)
pH, probably because the addition rate was inadequate to
alter soil chemistry. Furthermore, Luo et al. (2017) found
an increased microbial population in a sandy loam soil due
to the application of bamboo stick biochar. The high pH,
EC, and available K ­ + in biochar improved the soil proper-
ties required for microbial colonization (Chen et al. 2017a).
The elevated soil nutrient contents following biochar
application to soil could promote the activity/growth of soil
biota. For instance, application of maize straw-derived bio-
char at 30 t ha−1 to a silt loam soil increased the soil organic
C, total N, and total P, thus, increasing the availability of
soil nutrient (Luo et al. 2017). As a result, increased rela-
tive abundance of AMF and the AMF/saprotrophic fungi
ratio were observed when compared to the control (Luo et al.
2017).
In fact, the application doses of biochar and soil types
are important when considering the performance of soil
biota in biochar-amended soils. Abujabhah et al. (2018)
reported that biochar addition had a greater impact on
13
 Biochar
Fig. 4  SEM images of fungal hyphae grown on peanut shell biochar (pyrolyzed at 500 °C) surface (Images provided by the Hong Kong Univer-
sity of Science and Technology)
the bacterial diversity in three different-textured soils.
Furthermore, the results of that study suggested that soil
type and biochar rates affect the community structure and
the abundance of N-cycling bacteria in the soil (Abujab-
hah et al. 2018). Nevertheless, Luo et al. (2017) noted
that application of high quantities of biochar has greater
potential to increase fungal than bacterial diversity. Tre-
seder and Allen (2002) found that nutrient additions
by adding biochar promoted the growth of the AMF in
nutrient-poor soils. In addition, biochar alters N/P ratio
in soil and, therefore, the fungal root colonization rate
(Miller et al. 2002). Conversely, Nie et al. (2018) found
that the fungal population decreased with biochar applica-
tion, which may be attributed to increased P input through
biochar application (Bolan 1991).
13
3 Impact of biochar on soil microbial
activity and community structure
3.1 Soil respiration and organic carbon
mineralization
Organic amendments such as biochar and compost may
alter soil systems and organic C mineralization rate (Awad
et al. 2017; Cross and Sohi 2011). However, amendments
such as biochar increase recalcitrant organic carbon (OC)
compared to other forms of organic amendments, such as
plant residues, compost, and animal manure (Kimetu and
Lehmann 2010). Consequently, ­CO 2 evolution, the pri-
mary pathway by which ­CO 2 fixed by terrestrial plants
Biochar
returns to the atmosphere, can be controlled by adding
biochar to soil (Cross and Sohi 2011; Meier et al. 2017b).
Biochars produced by fast pyrolysis at low temperatures
may result in incomplete pyrolysis of biomass and those
biochars are more vulnerable to degradation (Bruun et al.
2012). Biochars produced at slow pyrolysis at high tem-
peratures result in complete pyrolysis of biomass due to
longer residence time. Thus, incompletely pyrolyzed bio-
chars have more labile C and the utilization of labile C by
soil microorganisms accelerates the biochar decomposi-
tion (Bruun et al. 2011). In contrast, completely pyrolyzed
biochars have more recalcitrant C and hence resistance to
microbial degradation (Bruun et al. 2011; Farrell et al.
2013).
There are many benefits of biochar applications, includ-
ing soil and water quality improvement, climate change
mitigation, energy production, and waste management;
however, initially, biochar application to the soil was
intended for long-term soil C sequestration (Ahmad et al.
2014; Lehmann et al. 2006a; Wong et al. 2017). Recently,
studies have highlighted the influences of biochar applica-
tion on soil OC mineralization. Junna et al. (2014) noticed
lower rates of OC mineralization for wheat straw biochar
produced at high temperature (600 °C) than at low tem-
perature (300 °C) and wheat straw itself. An interesting
finding in this study was that adding biochar decreased
wheat straw mineralization. However, this study did not
reveal actual mechanisms and processes involved in this
biochar-altered straw mineralization in the soil. Another
short-term soil incubation study found that microbial res-
piration as indicated by ­CO2 emission was increased by
adding switchgrass biochar pyrolyzed at 500 °C to the soil,
which was attributed to the increased amounts of read-
ily available C for microorganisms as a short-term energy
source (early stage of incubation) (Hawthorne et al. 2017;
Smith et  al. 2010). Interestingly, Pokharel and Chang
(Pokharel and Chang 2019) found that biochars produced
from manure pellet and woodchip have different effects on
­CO2 emission from bulk and rhizosphere soils. As bulk
soil alone may result in incorrect conclusions about the
impact of biochar on soil processes, care should be taken
when using different types of biochar in different soils.
Long-term biochar application may lead to enhanced C
storage in the soil as the complex aromatic and aliphatic
compounds are resistant to microbial degradation (Novak
et al. 2010; Smith et al. 2010). In relation to this, Xu et al.
(2018a) documented that there is a negative priming effect
and no obvious C mineralization in biochar-amended soils
in some cases. These reported findings imply that the
application of biochar would have altered microbial func-
tions and associated metabolisms of OC mineralization in
the biochar-amended soil, due to biochar-induced changes
in physicochemical properties of the soil.
Biochar interacts with native soil organic matter (SOM)
in a complex way; biochar may either have a positive or
negative priming effect or have no effect on native SOM.
This depends upon the feedstock type, pyrolysis temperature,
and organic matter level of soil. The soils richer in organic
matter status provide positive priming effect of native SOM
due to biochar addition and vice versa (Wang et al. 2016).
Biochar has high C sequestration potential and long-term
influence on native SOM.
One of the impacts of biochar on the soil is its influence
on bioavailable moisture dynamics in wetting–drying cycles
of the amended soil. Due to the high porosity and surface
area, the increased water availability may favor the sur-
vival of microbes during dry periods by providing a better
habitat, leading to enhanced microbial activities and res-
piration (Johnson et al. 2017; Junna et al. 2014). Reduced
microbial respiration was observed during the dry period,
owing to limited nutrients’ availability and the inactivation
or death of microorganisms (Junna et al. 2014). Conversely,
increased respiration upon rewetting was explained by the
greater availability of substrates/nutrients and reduced dif-
fusive limitations for substrates, resulting in the activation of
microorganisms and the subsequent utilization of the readily
available C (Junna et al. 2014; Rahman et al. 2018).
It is important to identify the biochar’s effect on gase-
ous losses of N from soil as there are many environmental
impacts due to the N ­ H3 and ­N2O emissions. For instance,
Thangarajan et al. (2018) recently examined the effect of
biochar and dicyandiamide (DCD, a chemical nitrification
inhibitor), on N transformation and N losses in soils treated
with organic amendments. The results showed 75% reduc-
tion in ­N2O emission by DCD treatment while increasing
­NH3 emission. However, biochar exhibited 23 and 43%
reduction in ­N2O and ­NH3 emissions from the soils, respec-
tively. These findings indicate the potential of biochar to
minimize N losses caused from ­NH3 volatilization and ­N2O
emission.
3.2 Soil enzyme activities
Many biogeochemical processes in the soil are facilitated
or catalyzed by soil microorganisms via secreted extracel-
lular and/or intracellular enzymes (Sabale et al. 2015). Soil
enzymes are important indicators of soil quality as they are
directly associated with microbial activity and biogeochemi-
cal cycling of nutrients (Ahmad et al. 2012; Das and Varma
2010; Hashimoto et al. 2009). Degraded soil quality may
be restored by biochar amendments through improving soil
biological properties, such as enhancing enzyme activity.
For instance, evaluation of changes in dehydrogenase activ-
ity in biochar-amended/unamended soils is a good indicator
for the evaluation of degree of recovery of degraded soils
(Demisie et al. 2014). Increased dehydrogenase activities in
13
 Biochar
biochar-amended soils were observed with different types
of biochar under different soil conditions (Bhaduri et al.
2016; Irfan et al. 2019; Mierzwa-Hersztek et al. 2016). The
enhanced dehydrogenase activities in biochar-treated soils
can be attributed to the addition of labile organic matter
and the high content on volatile matter of the used biochars
(Gascó et al. 2016).
A fluvo-aquic soil amended with biochar increased the
activity of some extracellular enzymes (such as ß-glucosi-
dase, ß-D-cellobiosidase, ß-xylosidase, and a-glucosidase,
and sulfatase) involved in soil C and sulfur (S) cycling
(Wang et al. 2015). However, this effect seemed inconsistent
and was depended on the rate of biochar addition. For exam-
ple, low rate of maize biochar application (0.5% [(w/w])
elevated the enzyme activity in the soil, whereas higher rates
of maize biochar application (1.0, 2.5, and 5.0% [w/w]) low-
ered the enzyme activity (Wang et al. 2015). Furthermore,
Demisie et al. (2014) found that dehydrogenase activity in
a degraded red soil was elevated by the application of oak
wood and bamboo biochar at 0.5% (w/w). In their study,
β-glucosidase activity was observed to be increased only
in the soil amended with bamboo biochar at 0.5 and 1.0%.
Moreover, urease activity was increased in soil amended
with oak wood biochar (0.5 and 2.0%) and bamboo biochar
(0.5%) at different application rates (Demisie et al. 2014).
However, it remains to be investigated as to how biochar reg-
ulates soil enzyme activities and what processes are involved
in this interaction.
Teutscherova et al. (2018) reported that liming effect of
biochar reduced Al toxicity in degraded acidic soils, which
was accompanied by increased dehydrogenase activities.
However, decreased β-glucosidase, β-glucosaminidase, and
phosphatase activities were observed for biochar-amended
Acrisol (Teutscherova et al. 2018). This can be attributed
to the adsorption of enzymes on biochar surfaces making
them unavailable for enzyme activities, for instance, > 99%
of β-glucosidase was adsorbed by chestnut wood biochar
surface (Lammirato et al. 2011). The study concluded that
low specific surface area and low porosity of biochar would
be helpful to maintain an active and protected enzyme pool
in soils (Lammirato et al. 2011). However, it is difficult to
draw a conclusion about causal relationship between bio-
chars and soil enzyme activities, due to the wide range
of biochars, soil conditions, and soil enzymes in the soil
environment (Table 1). The net effects of biochars on soil
enzyme activities can be indirect (e.g., microbial synthesis)
and direct (e.g., surface adsorption). Despite the beneficial
role of microorganism in the soil, soil-borne pathogens can
adversely affect plant growth. Jaiswal et al. (2018) revealed
that biochar could immobilize and deactivate enzymes which
degraded cell walls and detoxify metabolites produced by
Fusarium oxysporum f. sp. radicis lycopersici and, thus,
protect crop plants against soil pathogens. In general, these
13
studies showed that soil enzyme activity in biochar-amended
soils was dependent on the chemical composition of biochar,
biochar application rate, type of soil, and the nature of soil
enzymes (Fig. 1).
3.3 Soil microbial community structure
Soil microbial community structure in biochar-amended
soils has been studied with low, moderate-to-high-resolu-
tion techniques including PLFA, qPCR, DGGE, TGGE, and
DNA and RNA analyses (Fig. 5). A biochar-treated acid
soil was evaluated for soil microbial communities by PLFA;
where increased PLFA was observed compared to the con-
trol soil after 431 days of incubation (Luo et al. 2018). It has
been demonstrated that higher surface area of biochar could
adsorb water, nutrients, and soluble C which in turn facili-
tate the microbial colonization (Luo et al. 2018). Moreover,
the PLFA revealed that Gram-positive bacteria used biochar
C as substrate from the biochar produced at 350 °C than
that at 700 °C. Thus, the study suggested that the pyrolysis
temperature of biochar is an important factor controlling
the microbial community structure in soils regardless of the
soil pH (Luo et al. 2018). In contrast, several other studies
have observed a decrease in PLFA contents following the
biochar application into soils (Ameloot et al. 2014; Gomez
et al. 2014). As shown by Wang et al. (2015), the relative
abundances of fungi and bacteria, and the PLFA contents
were decreased with increasing application rate of maize
biochar. This can be explained by the alkalinity nature of
biochar that tends to inhibit soil microbial activity in the
short term and thereby reduce the community composition.
As soil pH is a key determinant of microbial community
structure and the evolution of certain functional groups
(Cheng et al. 2013; Lauber et al. 2009), it is not surprising
to have observed biochar amendments altered soil microbial
community structure and functions in the literature. Gomez
et al. (2014) noticed a reduction in PLFA concentrations
in three different soils with increasing application rate of
biochar. The authors cautioned that higher loading rates of
biochar could modify the soil structural properties and added
toxic compounds can negatively affect soil microorganisms.
However, this negative effect may also depend on the soil
condition. As revealed by Awad et al. (2018), application of
biochar reduced the total biomass measured using the PLFA
analysis at high redox potential, while increased total PLFA
biomass resulted in a low redox potential. The decreased
available C at higher redox potential and the increased pH
were responsible for these biomass changes (Awad et al.
2018).
Changes in microbial community composition can be
further studied using high-resolution techniques under dif-
ferent soil conditions using various biochar types. Biochar-
treated paddy soils were evaluated for bacterial and fungal
Biochar
Fig. 5  Methods involved in determining microbial community composition in biochar-amended soils and their relative effectiveness in terms of
resolution (modified from Nannipieri et al. (2003))
community structure and abundance using 16S rRNA and
18S rRNA gene, which were characterized using terminal-
restriction fragment length polymorphism (T-RFLP) com-
bined with clone library analysis, DGGE and qPCR assay
(Chen et al. 2013). The study found that gene copy num-
bers of bacterial 16S rRNA were increased, while that of
fungal 18S rRNA was decreased with biochar amendments.
Moreover, DGGE bands of fungal 18S rRNA gene revealed
that higher application rate (40 t ha−1) of biochar resulted
in inhibiting some bands associated with Ascomycota and
Glomeromycota. The study concluded that increased soil pH
and improved soil fertility might have supported to increase
the diversity of the bacterial community, whereas highly
stable organic C of biochar restricts the availability of sub-
strate for the fungi for their metabolism (Chen et al. 2013).
To identify the change of bacterial diversity and commu-
nity structure, DNA extraction, PCR amplification of 16S
rRNA gene fragments, and DNA sequencing were done in
a biochar-treated soil (Xu et al. 2016). The study revealed
that bacterial diversity increased with the increasing biochar
application rate. However, the relative abundance of some
phyla was decreased, while some others were increased.
These different observations in the above studies could be
attributed to several factors. Microorganisms which have
the ability to degrade recalcitrant C compounds are likely
to be predominant in biochar-amended soils (Kirby 2005;
O’neill et al. 2009). Moreover, acidophilic microorganism,
for example the bacteria phylum Acidobacteria, tends to
be diminished due to the increasing alkalinity with biochar
amendments (Mao et al. 2012). Nevertheless, increase of
soil moisture following the application of biochar tended to
decrease some microorganisms such as Gemmatimonadetes
as they prefer dry soil conditions (DeBruyn et al. 2011).
The above results of various studies have shown that
biochar has a potential to change the microbial community
structure in various degrees as a function of time, soil condi-
tion, and the type of biochar.
3.4 Response of soil microbes to biochar
amendment in contaminated soils
Many studies have demonstrated that biochar incorporation
into the soil can help to reduce the availability of various
contaminants in the soil. It is well known that biochar has
the capacity to immobilize organic/inorganic contaminants
in soils via various mechanisms such as complexation, sorp-
tion, precipitation, co-precipitation, and ion exchange. Those
processes alter soil chemical properties in biochar-amended
soils (Ahmad et al. 2014; Mohan et al. 2014). Yet, it has
been largely unknown if biochar improves or retards the bio-
logical activity in contaminated soils. Thus, it is important
to identify the response of soil microorganisms to biochar
amendments in contaminated soils, which is summarized
in Table 3.
Biochar may indirectly affect soil microbial commu-
nity structure and functions in heavy-metal-contaminated
soils, since heavy-metal toxicity may suppress or inhibit
soil microbial growth and alter soil microbial community
13
 Table 3  Effects of biochar application on soil microbial community/structure in contaminated soil

13
Biochar feedstock Production Dose (%) Soil type Contaminant (s) in soil Soil microbial response Possible mechanism (s) References
temperature involved
(°C)

Sugarcane bagasse
Wheat straw
Vegetable waste pine cone
Wood bark
Cocopeat
Palm kernel shell
Date palm waste
Poultry, cow, and sheep
manures
Gliricidia sepium wood
Gliricidia sepium wood
Chicken manure
Gliricidia sepium wood
Hardwood-derived
450 0.1–0.2 Sandy loam Cd, Cu, and Pb Actinomycetes popula- Reduced HMs bioavail- Nie et al. (2018)
tion: + 280% fungal popu- ability
lation: − 62%
450 0.7–2.9 Anthrosol Cd and Pb Actinomycetes population: Reduced HMs bioavail- Cui et al. (2013)
+19–38% fungi popula- ability
tion: +370–930%
200 0, 5 Sandy loam As and Pb Increased the microbial Immobilized HMs in soil Igalavithana et al. (2017a)
community abundance and
biogeochemical reactions
500 0, 5 Sandy loam Heavy metal No considerable effect on Immobilized HMs in soil Igalavithana et al. (2017b)
the microbial community
abundance and activity
300 0.5-3 NA Heavy metal Increased soil organic C, Immobilized HM in soil Al-Wabel et al. (2017)
soil respiration, and soil
microbial biomass C
450 5 NA Cr(VI) spiked Supported to reduce Cr(VI) Reduced Cr(VI) to Cr(III) Mandal et al. (2017)
to Cr(III)
NA 1-5 NA Heavy metal Increased soil microbial Reduced the bioavailable Seneviratne et al. (2017)
biomass C HM. Plant growth-pro-
moting bacteria increased
plant growth under HM
stress conditions
900 1-5 NA Cr Increased bacterial and Reduced the mobility, bio- Herath et al. (2017)
fungal counts availability, and phytotox-
icity of Cr in tomato plants
500 5–10 Sedimentary alfisol Cu Increased microbial activity Promoted the plant growth Meier et al. (2017a)
and dehydrogenase activity and reduced the Cu bio-
availability
300 and 700 1–5 NA Serpentine soils Fungal–bacterial co-inoc- Reduced heavy-metal toxic- Bandara et al. (2017)
ulated Biochar increased ity and assisted in plant
overall soil enzymatic growth
activity
NA 50 Silty sand As, Cd, Cu, Pb, and Zn Assisted in PAH degradation Significant reduction in PAH Beesley et al.( 2010)
concentration

NA not available, BC biochar, HM heavy metal, SOC soil organic carbon, PAH polyaromatic hydrocarbons
Biochar
Biochar
structure (Xu et al. 2018b). For instance, Hu et al. (2014)
revealed negative correlations between total and avail-
able heavy metals and microbial biomass carbon/nitro-
gen/MBN. Metal immobilization also contributes to the
reduction of bioavailable heavy metals by the presence
of ample O-containing functional groups on biochar sur-
faces (Abdelhafez et al. 2014). Nie et al. (2018) performed
a field experiment in a soil contaminated with Cu, Cd,
and Pb to examine the effects of biochar on availability
of heavy-metal and soil microbial activity. Application of
sugarcane bagasse-derived biochar at a rate of 3.0 t ha−1
increased soil bacterial population by 2.8 times, compared
to the control (Nie et al. 2018). However, the fungi popula-
tion was reduced, suggesting negative effects of biochar
application on fungal colonization. Prayogo et al. (2014)
observed that biochar amendment increased the popula-
tion of Gram-negative bacteria and actinomycetes in soil.
Chen et al. (2013) discovered that wheat straw biochar
diminished the fungal gene abundance while changing
community structure of fungi. The contrasting effects of
biochar on bacteria and fungi might be caused by the high
levels of mineral elements and the organic compounds in
biochar (Lehmann et al. 2011), and the increase in soil pH
(Chen et al. 2013). Furthermore, the sorption of organic C
and organically bound nutrients on biochar may hinder soil
microbial growth (Elzobair et al. 2016). Cui et al. (2013)
also reported that heavy metals (Pb and Cd) in soil alter
diversity, activities, and population size of microorgan-
isms. Biochar application alters soil pH and reduced the
heavy-metal bioavailability. It increased the population of
actinomycetes and fungi by 39 and 930%, respectively,
when compared with the control. A soil contaminated
with Cd and Pb reduces the total PLFA concentration to
a greater extent compared to the uncontaminated soils
indicating the metal toxicity on soil microorganism (Xu
et  al. 2018b). However, application of biochar allevi-
ated the metal toxicity in soil while increasing the PLFA
concentration which shows the reduced metal toxicity
and enhanced soil microbial community composition as
affected by biochar application. Application of vegetable
waste biochar produced at 200 °C improved the soil qual-
ity of heavy-metal-contaminated soils, through increas-
ing soil microbial abundance and associated microbial
activities which were proven by dehydrogenase activity
and fatty acid methyl ester (FAME) analysis (Igalavithana
et al. 2017a; Meier et al. 2017b). In contrast, Igalavithana
et al. (2017b) reported that the application of umbrella
tree (Maesopsis eminii)-derived biochar to a heavy-metal-
contaminated soil had no effect on microbial abundance
determined by FAME analysis and dehydrogenase activity
in a short term. The low surface area and low volatile mat-
ter content of the resultant biochars reduced the microbial
community and the activity in the soils amended with bio-
chars (Igalavithana et al. 2017b).
Elevated metal concentrations in soils can disrupt micro-
bial functions, resulting in adverse effects on the ecosys-
tem. Therefore, improving soil microbial performance with
amendments such as biochar would positively affect the
overall ecosystem health. Mandal et al. (2017) observed that
application of manure biochar enhanced Cr(VI) reduction in
Cr(VI)-spiked soil. Stimulation of the soil microbial com-
munity with the addition of biochar assisted the conversion
of Cr(VI) into less toxic Cr(III). This was mainly due to the
provision of C and protons to support microbial activities in
the soil (Ahmad et al. 2014; Choppala et al. 2012; Herath
et al. 2017; Mandal et al. 2017; Xia et al. 2019). Moreover,
chicken manure-derived biochar was used in a Cu-contam-
inated soil to reduce Cu bioavailability successfully and
increase microbial activity in the soil because of the storage
of Cu in metal-tolerant structures of the soil microbes (Meier
et al. 2017a). Ahmad et al. (2016) observed that biochar
produced at a low temperature could increase the abundance
of fungi, Gram-positive, and Gram-negative bacteria, actino-
bacteria, and actinomycetes in a Pb- and As-contaminated
soil. As a result, the capacity of biochar to change the soil
pH, organic matter content, and available nutrients has a
great impact on soil microbial activity in contaminated soils
amended with biochar (Beesley et al. 2010, 2011; Huang
et al. 2017).
A PCR-DGGE-based analysis was used to elucidate the
effect of biochar on indigenous microbial community com-
position in a heavy-metal-contaminated sediment (Liu et al.
2018). As reveled by the 16S rRNA sequencing analysis,
biochar application adversely affected the performance of
indigenous microorganism, resulting in reduced abundance
and diversity. Although the biochar caused to disappear some
bacteria species, the appearance of new bacteria species and
genera was observed. The study concluded that, although
the biochar caused to reduce the metal toxicity to the sedi-
ment microorganisms, biochar could modify the sediment’s
physicochemical properties, which may negatively affect the
indigenous microorganisms. Chen et al. (2017b) carried out
an experiment to assess the effect of biochar on diversity of
bacterial community in river sediment contaminated with
heavy metals. As seen in amplified 16S rDNA fragments in
DGGE profile, application of biochar decreased the number
of bands in DGGE profile at the early stage of the experi-
ment (1–5 days) indicating the reduced bacterial community.
However, the bands were significantly increased at the later
stages (7–50 days). The reduced bacterial communities at
the early stages can be attributed to the accelerated microbial
succession caused by biochar-induced microbial activities
and growth. As a result, the dominant microbes may occupy
the space and consume the nutrients rapidly which lead to
inhibit the other microorganism (Huang et al. 2017). On the
13
 Biochar
other hand, reduction of metal toxicity caused by biochar
and the changes in environmental conditions over the time
could account for the increased bacterial community at the
later stages (Chen et al. 2017b).
In addition to inorganic contaminants, organic contami-
nants such as polychlorinated biphenyl (PCB), polycy-
clic aromatic hydrocarbons (PAHs), organochlorides, and
nitroaromatic compounds are often found in soil environ-
ments. Biochar can also be used to remediate these organic
contaminated soils, resulting in their reduced toxicity, mobil-
ity, and bioavailability, which was attributed to microbial
activity primed/regulated by biochar application (Beesley
et al. 2010, 2011). Rhodes et al. (2010) observed a > 99%
reduction of phenanthrene in a contaminated soil with 0.1%
activated biochar amendment as biochar addition acceler-
ated microbial mineralization of phenanthrene and, thus,
its detoxification in the soil. In addition, microbial-assisted
degradation of chlorpyrifos and carbofuran was observed
in spring onion-cultivated soil amended with Eucalyptus
spp.-derived biochar (Yu et al. 2009). These effects may
be explained by the increased microbial biomass because
of increased supply of nutrients and available organic
compounds by biochar, which are readily available energy
sources for soil microorganisms. Other studies also found
that amendment of hardwood-derived biochar significantly
decreased PAH concentrations in the soil, resulting in > 50%
reduction of the heavier 4- and 5-ringed PAHs and a > 40%
reduction of the lighter 2- and 3-ringed PAHs, as compared
with the control (Beesley et al. 2010). In fact, the changed
physicochemical properties of biochar were responsible for
the reduction of a large proportion of PAHs in the soil. How-
ever, microbial degradation was also involved because of the
stimulation of microbial activity by biochar.
4 Conclusions and future research
directions
Biochar application has a significant potential for improving
soil health in different soil systems with or without exposure
to contamination by various heavy metals and/or organic
pollutants. The physicochemical properties of biochar such
as pH, EC, volatile organic compounds, surface and struc-
tural properties, and elemental composition play a major
role in influencing the soil microbial community structure.
Application of biochar into soils has been proven to enhance
soil microbial activity, diversity, and abundance. Especially,
provision of favorable microbial habitats and substrates for
their metabolic activities enhances microbial activities,
thereby promoting some ecologically important functions
such as enzyme activities, element cycling, decontamination
of polluted soils, and plant productivity. Depending on the
purpose of use, different types of biochar (i.e., feedstock type
13
and pyrolysis conditions) should be considered in promoting
soil health and productivity. With biochar as a potential soil
amendment for improving soil health by altering microbial
activities and functions, many aspects are still to be stud-
ied. Given the current knowledge on biochar and microbial
responses in soils discussed in this review, we recommend
the following areas for further research:
1. Most studies have evaluated the potential value of bio-
char application in improving the soil productivity in
relation to altering soil physicochemical properties. It
is also important to examine the value of this C rich
material in improving soil health through its effects on
microbial diversity and functions.
2. Depending on the biochar type as well as the soil type
and the composition of microorganisms, microbial inter-
actions with soil and plants may largely vary. Therefore,
studying the interactions of microorganisms with vari-
ous types of biochar, different biochar application rates,
and different plant species over the long term is required
to assess the magnitude of biochar effects on soil micro-
organisms over time, under different conditions.
3. Studies on biochar and microbial activities and interac-
tions in soils have so far been mainly focused on small-
scale laboratory incubation and greenhouse pot experi-
ments. It is recommended to conduct large-scale field
trials to investigate the long-term soil and plant interac-
tions with microbes as impacted by biochar application
with temporal changes in such long-term studies.
4. Current research on biochar and their effects on soil
microbial community structure, especially in contami-
nated soils, is mostly empirical. Biochars with different
physicochemical properties and application rates were
added to different soil types (soil texture and parent
material). The soil microbial community or enzymatic
activities were then measured after a certain period.
The major limitation of this approach is that the mecha-
nisms for the expected outcome are unknown. Further
research using biochar as a growth promotor of specific
soil microorganisms to achieve a desired goal (such as to
promote soil nutrient cycling) should use a customized
biochar (proactively select biochar feedstock and pro-
duction conditions) based on this review and the greater
literature.
5. Many studies have shown that remediation of inorganic/
organic contaminated soils with biochar application
is effective. Only a few studies have investigated the
changes in microbial community and activity in soils
after biochar application. It is important to understand
what specific microorganisms are involved in remedia-
tion of soils and the underlining mechanisms.
6. Biochar is often applied alone in soil as an amendment.
In fact, biochar possesses a distinct and highly porous
Biochar
structure with different pore sizes. Different soil micro-
organisms including bacteria and fungi can be inocu-
lated in biochar beforehand. It does not only increase the
colonization rate of desired groups of soil microbes, but
also speed up the presence of their beneficial effects in
biochar amended soils, especially for cleaning up soils
contaminated with different pollutants. Thus, the value
of biochar as a microbial carrier in crop production sys-
tems to improve soil health and productivity needs to be
examined.
7. It is difficult to isolate the impacts of biochar either
on a specific soil biological property or a specific soil
microbe in a microbial community. Sophisticated and
cutting-edge analytical techniques such as the fluo-
rescence in  situ hybridization (FISH) (Hubert et  al.
2016, Rice et al. 2016) and nanoscale secondary ion
mass spectrometry (NanoSIMs) can be adopted to help
advancing the scientific knowledge in this aspect.
8. Adopting high-resolution molecular based techniques
such as PLFA, PCR, DGGE, TGGE, and DNA and RNA
analyses is needed for detecting families, genera, or even
species level, which will be useful for better understand-
ing microbial community structure in biochar-amended
soils.
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