applied soil ecology 41 (2009) 50–58

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Charcoal production at kiln sites affects C and N dynamics

and associated soil microorganisms in Quercus spp.
temperate forests of central Mexico

Blanca Estela Gómez-Luna a,1, Ma. Cruz Rivera-Mosqueda a,1, Luc Dendooven b,2,
Gerardo Vázquez-Marrufo c,3, Vı́ctor Olalde-Portugal a,*
a
Laboratory of Ecological Biochemistry, Department Biotechnology and Biochemistry, Cinvestav Campus Guanajuato,
Km 9.6 Libramiento Norte Carretera Irapuato-León, Apdo. Postal 629, C.P. 36500, Irapuato, Guanajuato, Mexico
b
Laboratory of Soil Ecology, Department Biotechnology and Bioengineering, Cinvestav. Av. Instituto Politécnico Nacional 2508,
San Pedro Zacatenco, 07360 Ciudad de México, Mexico
c
Multidisciplinary Biotechnological Research Center, Veterinary and Animal Husbandry School, Universidad de San Nicolás de Hidalgo. Km 9.5
Carretera Morelia-Zinápecuaro, Apdo. Postal 53, C.P. 58262, Morelia, Michoacán, Mexico

article info abstract

Article history: Temperate forests dominated by Quercus spp. cover large parts of Central Mexico and rural
Received 20 May 2008 communities depend on these forests for wood and charcoal. The impacts of charcoal
Received in revised form production on selected chemical properties including C and N dynamics, and populations of
13 August 2008 ammonifiers, nitrifiers and denitrifiers were investigated on surface soils (0–15 cm) collected
Accepted 23 August 2008 during the dry and rainy season of these forests. Organic C was halved in soil at the kiln sites
compared to undisturbed forest soil. Concentrations of exchangeable Ca2+, K+ and Mg2+
increased >1.6 times at kiln sites and pH increased from 4.5 in undisturbed soil to 7.0 at kiln
Keywords: sites. The kiln sites had 1.3 times and 2.4 times lower microbial biomass C and N,
Ammonifiers respectively, than undisturbed forest sites during the rainy season. Although the effect
Charcoal production of charcoal production on NH4+, NO2 and NO3 concentrations was small, the ammonify-
Denitrifiers ing, nitrifying and denitrifiers were 16 times lower at the kiln sites than in the undisturbed
Nitrifiers forest soil. This research found that the charcoal production had a negative effect on the
Soil microbial biomass C and N cultivable microorganisms involved in N cycling and the soil microbial biomass C and N
compared to undisturbed forest soil. Differences in inorganic N dynamics were more
affected by seasonality, i.e. precipitation, than by charcoal production.
# 2008 Elsevier B.V. All rights reserved.

* Corresponding author. Tel.: +52 462 6239632; fax: +52 462 6245996.
E-mail addresses: bgomezl2000@yahoo.com.mx (B.E. Gómez-Luna), dendoove@cinvestav.mx (L. Dendooven),
hongo_indigo@prodigy.net.mx (G. Vázquez-Marrufo), v_olalde@yahoo.com.mx (V. Olalde-Portugal).
1
Tel.: +52 462 6239632; fax: +52 462 6245996.
2
Tel.: +52 555061 3311; fax: +52 555061 3313.
3
Tel.: +52 443 2958029; fax: +52 443 2958029.
0929-1393/$ – see front matter # 2008 Elsevier B.V. All rights reserved.
doi:10.1016/j.apsoil.2008.08.007
 applied soil ecology 41 (2009) 50–58
1. Introduction
In a forest ecosystem, soil is the dynamic entity regulating
most biogeochemical processes including nutrient storage
and recycling, organic matter decomposition, nitrogen fixa-
tion and emissions of greenhouse gasses, such as nitrous
oxide (N2O), methane (CH4) and carbon dioxide (CO2) (Mabu-
hay et al., 2003). Fire is a powerful controller of forest
ecosystems, causing changes in soil properties, as well as in
the distribution and productivity of plant communities
(Giovannini and Lucchesi, 1997; Hart et al., 2005). Fire also
exerts an indirect effect on soil conditions, due in part to post-
fire ash residues, and changes in microclimate. These factors
combine to affect C and N cycling through microbial
communities. Generally, significant changes in soil fertility
and soil microbial populations can be observed due to fire. The
effect of fire on the biogeochemical C and N cycling are
significant. Several studies have reported losses of soil
nutrients (C, N and P) due to fire, as well as changes in
nutrient availability (Kennard and Gholz, 2001; McIntosh et al.,
2005; Hatten et al., 2005). High temperature combustion of
organic matter (>200 8C) can result in volatilization and
potential loss of N present in organic complexes. Nitrogen
can also be lost through post-fire leaching (Giovannini and
Lucchesi, 1997; Choromanska and DeLuca, 2002; Mabuhay
et al., 2003; Johnson et al., 2005). Wild-fire combustion is often
incomplete, producing CH4, hydrocarbons and polyaromatic
compounds. Volatilization of organic compounds start at
100 8C, while carbonization takes place above 200 8C; lignin
breaks down >130 8C and hemicellulose >190 8C (Giovannini
and Lucchesi, 1997; González-Pérez et al., 2004). Temperatures
over 300 8C produce structural changes, humic and fluvic acid
decarboxylation, as well as an increase in aromatic com-
pounds that make charcoal recalcitrant to biological degrada-
tion (González-Pérez et al., 2004; Hart et al., 2005; Knicker et al.,
2005).
Fire also affects microorganisms, which can have negative
impacts in soil functioning as they control nutrient cycling and
organic matter formation and decomposition. Soil tempera-
tures >50 8C are enough to kill heat-sensitive microorganisms.
Lethal temperatures for fungi vary from 60 8C to 80 8C, while
bacteria normally resist heat better, being killed by tempera-
tures ranging from 100 8C to 120 8C (Choromanska and DeLuca,
2002; Hart et al., 2005). Microclimatical conditions change after
charcoal production. The vegetation cover disappears so that
the impact of rain is more direct resulting in erosion and
greater changes in soil water content, and an increase in solar
radiation so that temperature fluctuations are more extreme
(Fisher and Binkley, 2000; Hart et al., 2005). Most biological
processes are temperature and water dependent (Paul and
Clark, 1996). Increases in soil temperature will accelerate
organic matter degradation and nutrient cycling (Kaye and
Hart, 1998a,b) while drying and wetting affect soil microbial
activity (Paul and Clark, 1996).
Little information is available about the impacts of charcoal
in forest soils. Charcoal is still a commonly used fuel in many
rural communities of Latin America, Africa and the Middle
East (Manlay et al., 2000; Coomes and Burt, 2001; Glaser et al.,
2002; Vázquez-Marrufo et al., 2003; Nepstad et al., 2006).
Charcoal is made by burning wood in a kiln creating internal
51
suppressed combustion with the least possible flame. The
process lasts from 12 to 14 days (Vázquez-Marrufo et al., 2003).
During the production, a proportion of the biomass is
transformed to charcoal, which contains highly recalcitrant
molecules (Pietikäinen and Fritze, 1993; Zackrisson et al.,
1996). Losses of C and N during combustion can be as high as
68% (Zackrisson et al., 1996; Gardner and Whitlock, 2001;
Alexis et al., 2007; Eckmeier et al., 2007). Large amounts of CH4,
CO, CO2, alkanes, alkenes, oxygenated compounds and
particulate matter can be emitted during the process (Ezzati
et al., 2004; Bailis et al., 2005). In the central highlands of
Mexico charcoal is still produced in the Santa Rosa forest
(Guanajuato). Twenty kiln sites with a diameter of 7–10 m can
be found per ha so that up to 3% of the forest can get lost due to
charcoal production. Ex-kiln sites have different climate
conditions than other areas in the forest, such as higher solar
radiation, exposure to wind and water erosion, extreme
temperatures and a greater UV light incidence (Vázquez-
Marrufo et al., 2003). The soil is severely altered to a depth of
>20 cm at the kiln sites (Serrasolsas and Khanna, 1995; Prieto-
Fernández et al., 1998). The organic material is combusted
above 200 8C, but structural changes occur in the organic
material when the temperature is higher than 300 8C, e.g.
decarboxylation of the humic and fulvic acids and an increase
in aromatic components, making it far more resistant to
microbial degradation (González-Pérez et al., 2004; Hart et al.,
2005; Knicker et al., 2005). The microbial biomass composition,
number of bacteria and the basal respiration is reduced at the
kiln site (Zackrisson et al., 1996; Prieto-Fernández et al., 1998;
Pietikäinen et al., 2000; Glaser et al., 2002). Heating also
changes soil organic matter composition, enzyme activity, pH
and mineral N content (Guinto et al., 1999; Oguntunde et al.,
2004; Boerner et al., 2005; Guerrero et al., 2005). As such,
charcoal production has an immediate, but also a long-term
effect on the local environment (Zackrisson et al., 1996). All
vegetation is destroyed and reforestation can take years
erosion sets in further retarding the restoration of the
ecosystem. Even 15 years after the production of charcoal,
the soil has not recovered yet (Vázquez-Marrufo et al., 2003).
Currently, information about the impacts of charcoal produc-
tion on soil properties including nutrients, C and N dynamics
and soil microbial populations is lacking. Sustainable manage-
ment of ecosystems requires an understanding of the factors
that affect soil microbial populations, as there is a clear
relationship between microbial diversity, soil quality and
plant community (Mabuhay et al., 2003; Hart et al., 2005;
Vásquez-Murrieta et al., 2006). Thus, the objective of this paper
is to determine the impacts of charcoal production on C and N
dynamics, key soil nutrients and soil microbial populations
and mineralization during the dry and rainy seasons of the
Santa Rosa forest in Mexico.
2. Materials and methods
2.1. Study site location and soil characterization
The research area is located in the Santa Rosa in the state of
Guanajuato, Central Mexico (Northern Latitude 208500 5900 –
218550 0500 and Western longitude 1008590 2800 –1008330 0900 ). The
52 applied soil ecology 41 (2009) 50–58

altitude of the study area is 2660 m above sea level, with mean
annual temperature ranging from 12 8C to 18 8C and an
average rainfall of 1100 mm (CONABIO, 2007). The temperate
forest covers 1486 km2 and is dominated by Quercus rugosa
Neé, the main species used for charcoal production.
The kiln sites were prepared in 1999 and used for one year.
The kiln site of approximately 7 m in diameter was prepared
by clearing all vegetation, removing the organic residue or
litter, Oi and Oe horizon, from the surrounding area and
flattening the soil (Vázquez-Marrufo et al., 2003). The wood is
piled up in the centre of the kiln site, covered with the litter, lit
and then covered with soil. Some airing points at a high of
1.5 m are left uncovered and the incomplete combustion with
a minimum of flame production is maintained for 12–14 days.
The temperature of the covering soil varies between 200 8C
and 300 8C. The soil is charred or carbonized to a depth of
20 cm and some of the charcoal produced is left at the site as a
waste product.
Three different locations at a distance of 50–70 m were
identified within the same stand of approximately one ha.
Three different points were sampled at each treatment and
location: (i) non-disturbed soil that was never used to produce
charcoal (the CONTROL treatment), (ii) soil where charcoal
was produced for one year in 1999 (the KILN treatment), and
(iii) a transition point located between soil where charcoal was
produced and the control soil, previously identified and
monitored prior to this study (the TRANSITION ZONE
treatment). The soil sampled in the dry and rainy season.
As such, the number of soil samples obtained from each
treatment in the dry and rainy season was nine (n = 9), but only
soil sampled in the rainy season was used for characterization
(Tables 1 and 2).
The Oi and Oe horizons had been removed for the
preparation of the kiln site, but some humus or the Oa
horizon remained at the KILN treatment. A thick layer of litter
and humus was found in the TRANSITION ZONE and
CONTROL treatments. The soils in the area are classified as
haplic phaeozems (PHh) according to FAO/UNESCO (1988).
2.2. Soil sampling
Soil samples were taken from the three sites of each treatment
(kiln, transition zone and control) at 0–15 cm soil depth. Soil
samples were taken in July 2004 and May 2005 representing the
rainy and dry season, respectively. A metal cylinder with
diameter 10 cm and 15 cm long was used to sample five soil
columns at random at each sampling site. The five soil
samples of each sampling site and treatment were pooled,
2 mm sieved and characterized, producing a total of nine soil
samples, triplicate samples from each treatment.
2.3. Selected physical and chemical properties
Soil pH was measured in 1:2.5 soil–H2O suspension using a
glass electrode (Thomas, 1996). The organic matter C content
was determined as described by Pulido and Aguilar (1992).
Total N was measured by the Kjeldhal method (Bremner, 1996)
and the soil particle size distribution by Bouyoucos method
(Gee and Bauder, 1986). Extractable P was determined as
described by Olsen et al. (1954). The cationic interchangeable
macronutrients (Ca2+, Mg2+ and K+) were extracted with
ammonium acetate 1N at pH 7.0 before being quantified by
Atomic Absorption Spectrometry (Plenecassagne et al., 1997).
2.4. Microbial populations
The abundance of microorganisms was determined by the
most probable number method. Ten g serial samples were
diluted from 101 to 106 in triplicate. The MPN cultural
medium contained the following minerals [(g l1) K2HPO4
(0.25), MgSO47H2O (0.125), NaCl (0.125), FeC6H5O7nH2O (0.02)],
trace elements [(mg l1) MnSO4H2O (2.5), Fe2(SO4)3 (2.5),

Table 1 – Selected soil properties at the kiln sites (KILN treatment) and boundary sites (TRANSITION ZONE treatment)
compared to an undisturbed forest with Quercus spp. as the dominant species (CONTROL treatment) at Santa Rosa
(Guanajuato, Mexico) sampled in the rainy season
Treatment pHH2 O Organic C Total N Sand Clay Silt USDA soil texture
(g kg1 soil) (g kg1 soil) (g kg1 soil) (g kg1 soil) (g kg1 soil) classification

KILN 7.0 (0.2)a Ab 55.7 (0.7) B 5.1 (0.6) B 640 (30) A 110 (30) A 250 (40) A Sandy loam
TRANSITION ZONE 5.4 (0.2) B 64.4 (0.7) B 4.8 (0.6) B 700 (30) A 70 (20) A 230 (20) A Sandy loam
CONTROL 4.5 (0.1) C 114.0 (0.6) A 6.7 (0.2) A 730 (60) A 50 (10) A 220 (50) A Sandy loam
a
Value between parenthesis is the standard error of the estimate.
b
Values with the same letter are not significantly different between the treatments (P < 0.05) (n = 9).

Table 2 – Selected soil properties at the kiln sites (KILN treatment) and boundary sites (TRANSITION ZONE treatment)
compared to an undisturbed forest with Quercus spp. as the dominant species (CONTROL treatment) at Santa Rosa
(Guanajuato, Mexico) sampled in the rainy season
Treatment Extractable P (mg kg1 soil) Ca2+ (mg kg1 soil) K+ (mg kg1 soil) Mg2+ (mg kg1 soil)

KILN 162 b (30)a Bb 9955 (675) A 1733 (216) A 1244 (223) A

TRANSITION ZONE 303 b (89) B 6711 (1191) B 1156 (151) B 978 (233) A
CONTROL 660 (143) A 2267 (133) C 1067 (480) B 533 (133) A
a
Value between parenthesis is the standard error of the estimate.
b
Values with the same letter are not significantly different between the treatments (P < 0.05) (n = 9).
 applied soil ecology 41 (2009) 50–58
Na2MoO42H2O (0.05), Na2B4O710H2O (0.05), Co(NO3)26H2O
(0.05), CdSO43H2O (0.05), CuSO45H2O (0.05), ZnSO47H2O
(0.05), MnSO4H2O (0.05)] and a selective substrate for each
functional group, i.e. asparagine for ammonifiers (0.2 g l1);
(NH4)2SO4 (2 g l1) for nitrifiers and KNO3 (2 g l1) for deni-
trifiers. The cultural medium tubes used to quantify the
nitrifying and denitrifying organisms were incubated at 28 8C
for 7 days (Alexander, 1982).
The microbial biomass C was determined as described by
Vance et al. (1987) and microbial biomass N as described by
Joergensen and Brookes (1990). Ethanol-free chloroform was
used to fumigate the soil (Mueller et al., 1992). The microbial
biomass C was calculated as [2.64  (K2SO4-extractable
organic C in fumigated soil  K2SO4-extractable organic C in
unfumigated soil)] (Vance et al., 1987) and the microbial
biomass N as [2.22  (K2SO4-extractable total N in fumigated
soil  K2SO4-extractable total N in unfumigated soil)] (Jenkin-
son, 1988).
2.5. Aerobic incubation
Twelve sub-samples of 25 g soil from each sampling location
were added to 120 ml glass flasks. Three were chosen at
random from each treatment and soil was extracted for
inorganic N with 100 ml 0.5 M potassium sulphate (K2SO4). The
samples were shaken for 60 min and filtered through What-
man No 42 paper1 and stored at 20 8C before analysis. This
provided zero-time samples. The remaining glass flasks were
placed in 945 ml glass jars containing one vessel with 10 ml
distilled H2O and one beaker with 20 ml 1 M NaOH to trap CO2
evolved. The jars were sealed and stored in the dark for 28 days
at 22  2 8C. An additional nine jars with one beaker of 10 ml
distilled H2O and one beaker of 20 ml 1 M NaOH were sealed
and served as controls to account for the CO2 trapped from the
air. After 7, 14 and 28 days, three jars selected randomly from
each treatment were opened and the vessel containing NaOH
was removed. A 5 ml NaOH aliquot was analyzed for CO2
trapped. The soil was removed from three flasks and extracted
for inorganic N as described for zero-time samples. All
remaining flasks were opened, aired for 10 min to avoid
anaerobicity, sealed and stored in the dark.
2.6. Statistical analyses
Concentration of NH4+, NO2, NO3, and microbial biomass C
and N were subjected to one-way analysis of variance using
Table 3 – Microbial biomass C and biomass N (mg kgS1 soil) in soil at the kiln sites (KILN treatment) and boundary sites
(TRANSITION ZONE treatment) compared to an undisturbed forest with Quercus spp. as the dominant species (CONTROL
treatment) at Santa Rosa (Guanajuato, Mexico) sampled in the rainy or the dry season
Treatment Microbial biomass C
1
Rainy (mg kg soil) Dry (mg kg
KILN 2177 (44)a Bb
TRANSITION ZONE 2969 (77) A
CONTROL 2724 (90) A
a
Values between parenthesis are standard errors of the estimate.
b
Values with the same letter are not significantly different between the treatments (P < 0.05) (n = 9).
53
PROC GLM (SAS Institute, 1989) to test for significant
differences between treatments with the Tukey’s Studentized
Range test. Significant differences between treatments for CO2
emissions were determined using PROC MIXED (SAS Institute,
1989). All data presented were the mean of three soil samples
and three plots, i.e. n = 9.
3. Results
3.1. Physical and chemical soil properties
Physical and chemical soil characteristics were affected by
charcoal production at the kiln site. Soil pH showed this trend:
kiln > transition zone > control (Table 1). The organic C and
total N contents of soil were lower in the TRANSITION ZONE
and KILN treatments than in the CONTROL treatment
(P < 0.05).
The Ca2+ content was intermediate in the TRANSITION
ZONE sites compared to the CONTROL and KILN treatments
(Table 2). Extractable P showed this trend: control > transition
zone > kiln. The K+ content was 1.5 times higher in the KILN
treatment compared to the TRANSITION ZONE and CONTROL
treatment (P < 0.05). The Mg2+ content was not affected by
treatment.
3.2. Microbial biomass C and N
The microbial biomass C was lower in the KILN treatment
compared to the TRANSITION ZONE and CONTROL treatments
in the rainy and dry season (P < 0.05) (Table 3). The microbial
biomass N was lower in the KILN treatment compared to the
TRANSITION ZONE treatment in the rainy and dry season
(P < 0.05). The microbial biomass N was lower in the
TRANSITION ZONE treatment compared to the CONTROL
treatment in the rainy and dry season (P < 0.05). Although
similar trends were found in the microbial biomass for the
rainy and dry seasons, there were greater differences in
microbial biomass between the transition zone and control
sites during the dry season.
3.3. Ammonifiers, nitrifiers and denitrifiers
The number of ammonifiers was lower in the KILN treatment
than in the CONTROL treatment in the rainy and dry season
(P < 0.05) (Table 4). The amount of NH4+ and NO2 oxidizers
Microbial biomass N
1
soil) Rainy (mg kg1 soil) Dry (mg kg1 soil)
406 (50) B 220 (10) C 52 (6) C
800 (54) A 476 (17) B 125 (10) B
990 (77) A 591 (35) A 213 (16) A
54 applied soil ecology 41 (2009) 50–58

Table 4 – Ammonifiers, NH4+ and NO2S oxidizers and denitrifiers in soil at the kiln sites (KILN treatment) and boundary
sites (TRANSITION ZONE treatment) compared to an undisturbed forest with Quercus spp. as the dominant species
(CONTROL treatment) at Santa Rosa (Guanajuato, Mexico) sampled in the rainy or the dry season
Treatment Ammonifiers NH4+ oxidizers NO2 oxidizers Denitrifiers

Rainy Dry Rainy Dry Rainy Dry Rainy Dry

(MPNa  103 (MPNa  103 (MPNa  103 (MPNa  103 (MPNa  103 (MPNa  103 (MPN g1 (MPN g1
g1 soil) g1 soil) g1 soil) g1 soil) g1 soil) g1 soil) soil) soil)

KILN 6 Bb 11 B 5B 20 B 7C 3C 147 B 49 B
TRANSITION ZONE 25 B 540 A 9B 77 B 34 B 38 B 97  103 A 140  103 A
CONTROL 970 A 970 A 230 A 518 A 110 A 110 A 140  103 A 140  103 A
a
MPN: Most probable number.
b
Values with the same letter are not significantly different between the treatments (P < 0.05) (n = 9).

was lower in the KILN and TRANSITION ZONE treatments than
in the CONTROL treatment in both the rainy and dry season
(P < 0.05). The amount of denitrifiers was lower in the KILN
treatment than in the TRANSITION ZONE and CONTROL
treatments in both the rainy and dry season (P < 0.05).
3.4. Dynamics of C and N
The emission of CO2 was linear in the soil sampled in the rainy
season, but slowed down after 7 or 14 days when sampled in the
dry season (Fig. 1a). The lowest C mineralization was found in
the KILN treatment. The concentration of NH4+ remained
constant in soil sampled in the rainy season, but increased
when soil was sampled in the dry season (P < 0.05) (Fig. 1b). The
concentration of NO2 remained <0.3 mg N kg1 soil and was
not affected by treatment or season (Fig. 1c). The concentration
of NO3 did not change over time when soil was sampled in
the dry season, but did increase when sampled in the rainy
season except in the CONTROL treatment (P < 0.05).
After 28 days, the amount of soil organic matter miner-
alized to N, i.e. the amount of inorganic N (NH4+ + NO2 +
NO3) at the end of the incubation minus the inorganic N at the
onset of the incubation, was 0.28% in soil at the KILN
treatment, 0.18% in the TRANSITION ZONE and 0.10% in the
CONTROL soil in the dry season and 0.18%, 0.13% and 0.01% in
the rainy season, respectively.

Fig. 1 – (a) Emission of CO2 (mg C kgS1 soil) and concentrations of (b) NH4+, (c) NO2S and (d) NO3S in soil where charcoal was
produced in the dry (*, KILN-dry) and rainy season (*, KILN-rain), at the transition zone of the charcoal production site in
the dry season (& TRANSITION ZONE-dry) and the rainy season (&, TRANSITION ZONE-rain) and in the forest soil in the dry
(^, CONTROL-dry) and rainy season (^, CONTROL-rain). Soil was incubated aerobically at 22 W 2 8C for 28 days. Bars are
Wone standard deviations.
 applied soil ecology 41 (2009) 50–58
4. Discussion
Charcoal production induced significant changes in the
selected soil properties studied at the kiln sites compared to
the undisturbed (CONTROL) sites. Increased pH at the kiln
sites can be explained by the accumulation of base-forming
cations (Ca2+, Mg2+, K+), production of alkaline primary oxides,
carbonates and the loss of organic acids (Ulery et al., 1993;
Giovannini and Lucchesi, 1997; Glaser et al., 2002; Pitman,
2006). Oguntunde et al. (2004) reported similar increases in pH
from 5.8 to 7.6 at a charcoal production site compared to
adjacent field soils in Ejura, Ghana. The concentrations of Ca2+
increased >4 times, Mg2+ >2 times and K+ >1.5 times
explaining the increase in pH at the kiln sites. Available P
decreased >4 times at the kiln sites compared to the
surrounding forest. Oguntunde et al. (2004) reported relative
increases of up to 329% for K+, 49% for Ca2+, 14% for Mg2+ and
169% for Na+ at a charcoal production site compared to
adjacent field soils. Losses of available P at the kiln sites might
be due to leaching, volatilization or ash losses (Leinweber
et al., 1999; Johnson et al., 2005; Murphy et al., 2006a,b).
However, Giovannini and Lucchesi (1997) and Kennard and
Gholz (2001) observed a post-fire increase of available P and
postulated that changes in P mineralization rates might
increase available P.
It is also well known that deforestation and litter removal
can have significant effects on soil organic matter content and
often more than halves it (Solomon et al., 2000). Charcoal
production also reduces soil organic matter and Oguntunde
et al. (2004) reported a 10% decrease in soil organic C at a
charcoal site compared to the undisturbed sites. In agreement
with previous findings, removal of the litter layer and
combustion halved organic C in the KILN treatment compared
the CONTROL treatment.
Similar to organic matter trends, microbial biomass C and
N in soil were reduced at the kiln sites in both the rainy and dry
season, which agrees with previous studies (Serrasolsas and
Khanna, 1995; Prieto-Fernández et al., 1998). The decreases in
microbial biomass C and N, however, were more pronounced
during the dry season than in the rainy season. The O and A
horizon were maintained in the CONTROL treatment, which
would reduce water losses and extreme temperatures in the
dry season. The soil in the KILN treatment lacked the O and A
horizon so was exposed to extreme temperatures and water
losses. The more extreme conditions in the KILN treatment
site explain the reductions observed in soil microbial biomass
C compared to the CONTROL treatment. The positive effects of
water availability, however, explain the less pronounced
effects of charcoal production in the rainy season. Prasad
et al. (1994) observed that converting forest to grassland or
arable land affected both the quantity and quality of microbial
biomass. It is well known that soil microorganisms depend on
soil organic C as substrate and decreases in soil organic matter
will decrease the soil microbial biomass. The microbial
biomass N as a percentage of soil organic N was higher in
the CONTROL treatment than in the KILN treatment, but not
the microbial biomass C as a percentage of soil organic C in the
rainy season. The microbial biomass C was 3.9% of soil organic
C in the KILN treatment in the rainy season and 2.4% in the
surrounding forest soil. In the dry season, it was 0.7 and 0.9,
55
respectively. The microbial biomass N was 4.3% of soil organic
N, in the KILN treatment in rainy season and 8.8% in the
surrounding forest soil. In the dry season, it was 1.0% and 3.2%,
respectively. Ross and Tate (1993) reported that the microbial
biomass C as a percentage of soil organic carbon in a forest soil
ranged from 1 to 3% and microbial biomass N was 5–9% of total
N, while Prasad et al. (1994) reported values of 2–3% for
microbial biomass C and 2–4% for microbial biomass N. It can
be speculated that the remaining organic material in the KILN
treatment was recalcitrant and did not provide the same
amount of easily decomposable C substrate as was found in
the forest soil and as such maintained lower amounts of
microbial biomass. Climatic conditions also had a large impact
on soil microorganisms. The microbial biomass C and N as a
percentage of soil organic C was lower in the dry than in the
rainy season. Murphy et al. (2007) also reported seasonal
changes in C and N pools in soils under arable and ley-arable
rotations in temperate environments. Moisture availability
affects microbial activity, nutrient cycling, regeneration of
forest vegetation and the functioning of ecosystems (Saynes
et al., 2005).
The microbial biomass C:N ratio was 9.9 in the KILN
treatment, 6.2 in the TRANSITION ZONE treatment and 4.6 in
the CONTROL treatment in the rainy season and 7.8, 6.4 and
4.6, respectively, in the dry season. There have been
suggestions that a higher microbial biomass C:N ratio, as in
the kiln treatment, may indicate more fungi with respect to
bacteria (Moore et al., 2000; Acosta-Martı́nez et al., 2004).
Although bacteria resist better the direct impact of heat than
fungi (Guerrero et al., 2005), the loss of easily decomposable
organic material due to charring and the formation of
recalcitrant organic material may favor fungi in the long
run. Sampling time, i.e. dry versus rainy season, appeared to
have little effect on biomass C:N ratio. In fact, microorgan-
isms involved in the N dynamics in soil, i.e. ammonifiers,
NH4+ oxidizers, NO2 oxidizers and denitrifiers, were lower in
soil at the kiln sites than in the surrounding forest soil. An
initial decrease in number of microorganisms commonly
observed after burning, is caused mainly by the effect of heat
liberated in the combustion. Burning causes severe heating of
soil, and this immediately kills some of the soil microbes
(Pietikäinen and Fritze, 1995). Some groups of soil micro-
organisms are more sensitive to heat than others especially
those that are highly specialized (Hart et al., 2005). For
instance, nitrite oxidizers, such as Nitrobacter spp. have a 99%
mortality rate at 80 8C wet soils and a 90% in dry soils (Hart
et al., 2005) and nitrifiers are very sensible to heating
(Serrasolsas and Khanna, 1995).
The emission of CO2 showed a flush in the dry season, but
not in the rainy season. It can be speculated that organic
material accumulated in soil in the dry season. Upon wetting
the organic material, especially the easily decomposable part,
will be mineralized causing a flush in emissions of CO2
(Updegraff et al., 1995). Less organic material will accumulate
in the rainy season as microbial activity is less inhibited so no
flush in C mineralization will be observed. Burning reduced
the amount of CO2 evolved after 28 days compared to the other
treatments. Burning reduces the amount of organic matter
incorporated in soil, soil organic C and thus C substrate for soil
microorganisms (Hamman et al., 2007).
56 applied soil ecology 41 (2009) 50–58
The amount of soil organic C evolved as CO2 within 7 days
was 0.56% at the kiln sites, 0.52% in the transition zone and
0.20% in the surrounding forest soil in the dry season and
0.34%, 0.23% and 0.19% in the rainy season, respectively.
Different explanations are possible to explain this phenom-
enon. First, the microbial efficiency for C was lower at the kiln
sites than in the surrounding forest soil. It has been reported
that the efficiency for recalcitrant organic material is lower
than for easily decomposable organic material (Van Veen
et al., 1985). Second, microbial biomass killed by fire or soil
organic matter became available as C substrate inducing an
increase in emission of CO2. Increases in easily decomposable
organic C after fire have been reported (Guerrero et al., 2005).
However, it can be assumed that this easily decomposable
organic material will be mineralized rapidly and will have no
effect on CO2 emissions after more than five years.
Specific qCO2 or the respiratory activity defined as the rate
of base respiration expressed per unit soil microbial biomass
(Anderson, 1994) is often used as an indicator of the energy
efficiency of soil microorganisms (Odum, 1985; Insam, 2001).
qCO2 calculated from the CO2 evolved between day 7 and 14,
i.e. basal respiration, and the soil microbial biomass was >2
times higher in the KILN treatment than in the CONTROL
treatment. A higher qCO2 can be interpreted as an indicator of
metabolic stress for the soil microorganisms (Ross and Tate,
1993; Prasad et al., 1994; Changming and Moncrieff, 2005). The
higher qCO2 value in the KILN treatment could be indicative of
a microbial community less efficient in assimilating the C
metabolized or an increase in their maintenance energy
requirement (Giller et al., 1998). The efficiency for C is lower
when the soil organic C is more resistant to degradation, i.e.
the proportion of recalcitrant soil organic matter was higher at
the kiln site, and the qCO2 was thus higher (Van Veen et al.,
1985).
Although the number of ammonifiers and nitrifiers was
significantly reduced by fire, burning had no effect on
ammonification or oxidation of NH4+ and NO2 according to
the concentrations of NH4+, NO2 and NO3. This may be
because burning is known to have a profound effect on soil
characteristics, but some of the effects are only short-lived
(Ford and Johnson, 2006). Additionally, the microorganisms
that were not killed might have increased their activity due to
the additional substrates available from dead microorgan-
isms. Guinto et al. (1999) reported that in a dry eucalypt forest
of south-east Queensland (Australia) extractable N and net N
mineralization were not affected by burning, but they
decreased after burning in a wet eucalypt forest.
5. Conclusions
Assessment of selected soil properties important for soil
functioning as affected by charcoal production is very crucial
for management decisions at the Santa Rosa Quercus spp.
temperate forest in central Mexico, where charcoal is
produced by rural communities as an energy source or to sell
at local markets. For example, this study found lower soil
organic C and higher Ca2+, K+ and Mg2+ and pH at the kiln sites
compared to the undisturbed forest soil. Charcoal production
had a negative effect on the cultivable microorganisms
involved in N cycle, i.e. ammonifiers, nitrifiers and denitrifiers,
and soil microbial biomass C and N. Differences in dynamics of
inorganic N in soil, however, were more affected by season, i.e.
dry or rainy season, than by charcoal production. The
deterioration in soil characteristics at the charcoal sites
compared to the undisturbed forest, including soil microbial
properties, should be considered so as to modify charcoal
production and preserve the soil.
Acknowledgements
We thank Rosalinda Serrato-Flores, Juan T Frı́as-Hernández and
L Antonio Jiménez-Magdaleno for technical assistance. This
project was funded by CONACYT-SEMARNAT project 2002-C01-
0129 (México) and CONAFOR project 2004-C04-01 (México).
B.E.G.-L. received grant aided support from CONACYT.
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