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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 51

1. Introduction suppressed combustion with the least possible flame. The


process lasts from 12 to 14 days (Vázquez-Marrufo et al., 2003).
In a forest ecosystem, soil is the dynamic entity regulating During the production, a proportion of the biomass is
most biogeochemical processes including nutrient storage transformed to charcoal, which contains highly recalcitrant
and recycling, organic matter decomposition, nitrogen fixa- molecules (Pietikäinen and Fritze, 1993; Zackrisson et al.,
tion and emissions of greenhouse gasses, such as nitrous 1996). Losses of C and N during combustion can be as high as
oxide (N2O), methane (CH4) and carbon dioxide (CO2) (Mabu- 68% (Zackrisson et al., 1996; Gardner and Whitlock, 2001;
hay et al., 2003). Fire is a powerful controller of forest Alexis et al., 2007; Eckmeier et al., 2007). Large amounts of CH4,
ecosystems, causing changes in soil properties, as well as in CO, CO2, alkanes, alkenes, oxygenated compounds and
the distribution and productivity of plant communities particulate matter can be emitted during the process (Ezzati
(Giovannini and Lucchesi, 1997; Hart et al., 2005). Fire also et al., 2004; Bailis et al., 2005). In the central highlands of
exerts an indirect effect on soil conditions, due in part to post- Mexico charcoal is still produced in the Santa Rosa forest
fire ash residues, and changes in microclimate. These factors (Guanajuato). Twenty kiln sites with a diameter of 7–10 m can
combine to affect C and N cycling through microbial be found per ha so that up to 3% of the forest can get lost due to
communities. Generally, significant changes in soil fertility charcoal production. Ex-kiln sites have different climate
and soil microbial populations can be observed due to fire. The conditions than other areas in the forest, such as higher solar
effect of fire on the biogeochemical C and N cycling are radiation, exposure to wind and water erosion, extreme
significant. Several studies have reported losses of soil temperatures and a greater UV light incidence (Vázquez-
nutrients (C, N and P) due to fire, as well as changes in Marrufo et al., 2003). The soil is severely altered to a depth of
nutrient availability (Kennard and Gholz, 2001; McIntosh et al., >20 cm at the kiln sites (Serrasolsas and Khanna, 1995; Prieto-
2005; Hatten et al., 2005). High temperature combustion of Fernández et al., 1998). The organic material is combusted
organic matter (>200 8C) can result in volatilization and above 200 8C, but structural changes occur in the organic
potential loss of N present in organic complexes. Nitrogen material when the temperature is higher than 300 8C, e.g.
can also be lost through post-fire leaching (Giovannini and decarboxylation of the humic and fulvic acids and an increase
Lucchesi, 1997; Choromanska and DeLuca, 2002; Mabuhay in aromatic components, making it far more resistant to
et al., 2003; Johnson et al., 2005). Wild-fire combustion is often microbial degradation (González-Pérez et al., 2004; Hart et al.,
incomplete, producing CH4, hydrocarbons and polyaromatic 2005; Knicker et al., 2005). The microbial biomass composition,
compounds. Volatilization of organic compounds start at number of bacteria and the basal respiration is reduced at the
100 8C, while carbonization takes place above 200 8C; lignin kiln site (Zackrisson et al., 1996; Prieto-Fernández et al., 1998;
breaks down >130 8C and hemicellulose >190 8C (Giovannini Pietikäinen et al., 2000; Glaser et al., 2002). Heating also
and Lucchesi, 1997; González-Pérez et al., 2004). Temperatures changes soil organic matter composition, enzyme activity, pH
over 300 8C produce structural changes, humic and fluvic acid and mineral N content (Guinto et al., 1999; Oguntunde et al.,
decarboxylation, as well as an increase in aromatic com- 2004; Boerner et al., 2005; Guerrero et al., 2005). As such,
pounds that make charcoal recalcitrant to biological degrada- charcoal production has an immediate, but also a long-term
tion (González-Pérez et al., 2004; Hart et al., 2005; Knicker et al., effect on the local environment (Zackrisson et al., 1996). All
2005). vegetation is destroyed and reforestation can take years
Fire also affects microorganisms, which can have negative erosion sets in further retarding the restoration of the
impacts in soil functioning as they control nutrient cycling and ecosystem. Even 15 years after the production of charcoal,
organic matter formation and decomposition. Soil tempera- the soil has not recovered yet (Vázquez-Marrufo et al., 2003).
tures >50 8C are enough to kill heat-sensitive microorganisms. Currently, information about the impacts of charcoal produc-
Lethal temperatures for fungi vary from 60 8C to 80 8C, while tion on soil properties including nutrients, C and N dynamics
bacteria normally resist heat better, being killed by tempera- and soil microbial populations is lacking. Sustainable manage-
tures ranging from 100 8C to 120 8C (Choromanska and DeLuca, ment of ecosystems requires an understanding of the factors
2002; Hart et al., 2005). Microclimatical conditions change after that affect soil microbial populations, as there is a clear
charcoal production. The vegetation cover disappears so that relationship between microbial diversity, soil quality and
the impact of rain is more direct resulting in erosion and plant community (Mabuhay et al., 2003; Hart et al., 2005;
greater changes in soil water content, and an increase in solar Vásquez-Murrieta et al., 2006). Thus, the objective of this paper
radiation so that temperature fluctuations are more extreme is to determine the impacts of charcoal production on C and N
(Fisher and Binkley, 2000; Hart et al., 2005). Most biological dynamics, key soil nutrients and soil microbial populations
processes are temperature and water dependent (Paul and and mineralization during the dry and rainy seasons of the
Clark, 1996). Increases in soil temperature will accelerate Santa Rosa forest in Mexico.
organic matter degradation and nutrient cycling (Kaye and
Hart, 1998a,b) while drying and wetting affect soil microbial
activity (Paul and Clark, 1996). 2. Materials and methods
Little information is available about the impacts of charcoal
in forest soils. Charcoal is still a commonly used fuel in many 2.1. Study site location and soil characterization
rural communities of Latin America, Africa and the Middle
East (Manlay et al., 2000; Coomes and Burt, 2001; Glaser et al., The research area is located in the Santa Rosa in the state of
2002; Vázquez-Marrufo et al., 2003; Nepstad et al., 2006). Guanajuato, Central Mexico (Northern Latitude 208500 5900 –
Charcoal is made by burning wood in a kiln creating internal 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 CONTROL treatments. The soils in the area are classified as
annual temperature ranging from 12 8C to 18 8C and an haplic phaeozems (PHh) according to FAO/UNESCO (1988).
average rainfall of 1100 mm (CONABIO, 2007). The temperate
forest covers 1486 km2 and is dominated by Quercus rugosa 2.2. Soil sampling
Neé, the main species used for charcoal production.
The kiln sites were prepared in 1999 and used for one year. Soil samples were taken from the three sites of each treatment
The kiln site of approximately 7 m in diameter was prepared (kiln, transition zone and control) at 0–15 cm soil depth. Soil
by clearing all vegetation, removing the organic residue or samples were taken in July 2004 and May 2005 representing the
litter, Oi and Oe horizon, from the surrounding area and rainy and dry season, respectively. A metal cylinder with
flattening the soil (Vázquez-Marrufo et al., 2003). The wood is diameter 10 cm and 15 cm long was used to sample five soil
piled up in the centre of the kiln site, covered with the litter, lit columns at random at each sampling site. The five soil
and then covered with soil. Some airing points at a high of samples of each sampling site and treatment were pooled,
1.5 m are left uncovered and the incomplete combustion with 2 mm sieved and characterized, producing a total of nine soil
a minimum of flame production is maintained for 12–14 days. samples, triplicate samples from each treatment.
The temperature of the covering soil varies between 200 8C
and 300 8C. The soil is charred or carbonized to a depth of 2.3. Selected physical and chemical properties
20 cm and some of the charcoal produced is left at the site as a
waste product. Soil pH was measured in 1:2.5 soil–H2O suspension using a
Three different locations at a distance of 50–70 m were glass electrode (Thomas, 1996). The organic matter C content
identified within the same stand of approximately one ha. was determined as described by Pulido and Aguilar (1992).
Three different points were sampled at each treatment and Total N was measured by the Kjeldhal method (Bremner, 1996)
location: (i) non-disturbed soil that was never used to produce and the soil particle size distribution by Bouyoucos method
charcoal (the CONTROL treatment), (ii) soil where charcoal (Gee and Bauder, 1986). Extractable P was determined as
was produced for one year in 1999 (the KILN treatment), and described by Olsen et al. (1954). The cationic interchangeable
(iii) a transition point located between soil where charcoal was macronutrients (Ca2+, Mg2+ and K+) were extracted with
produced and the control soil, previously identified and ammonium acetate 1N at pH 7.0 before being quantified by
monitored prior to this study (the TRANSITION ZONE Atomic Absorption Spectrometry (Plenecassagne et al., 1997).
treatment). The soil sampled in the dry and rainy season.
As such, the number of soil samples obtained from each 2.4. Microbial populations
treatment in the dry and rainy season was nine (n = 9), but only
soil sampled in the rainy season was used for characterization The abundance of microorganisms was determined by the
(Tables 1 and 2). most probable number method. Ten g serial samples were
The Oi and Oe horizons had been removed for the diluted from 101 to 106 in triplicate. The MPN cultural
preparation of the kiln site, but some humus or the Oa medium contained the following minerals [(g l1) K2HPO4
horizon remained at the KILN treatment. A thick layer of litter (0.25), MgSO47H2O (0.125), NaCl (0.125), FeC6H5O7nH2O (0.02)],
and humus was found in the TRANSITION ZONE and 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 53

Na2MoO42H2O (0.05), Na2B4O710H2O (0.05), Co(NO3)26H2O PROC GLM (SAS Institute, 1989) to test for significant
(0.05), CdSO43H2O (0.05), CuSO45H2O (0.05), ZnSO47H2O differences between treatments with the Tukey’s Studentized
(0.05), MnSO4H2O (0.05)] and a selective substrate for each Range test. Significant differences between treatments for CO2
functional group, i.e. asparagine for ammonifiers (0.2 g l1); emissions were determined using PROC MIXED (SAS Institute,
(NH4)2SO4 (2 g l1) for nitrifiers and KNO3 (2 g l1) for deni- 1989). All data presented were the mean of three soil samples
trifiers. The cultural medium tubes used to quantify the and three plots, i.e. n = 9.
nitrifying and denitrifying organisms were incubated at 28 8C
for 7 days (Alexander, 1982).
The microbial biomass C was determined as described by 3. Results
Vance et al. (1987) and microbial biomass N as described by
Joergensen and Brookes (1990). Ethanol-free chloroform was 3.1. Physical and chemical soil properties
used to fumigate the soil (Mueller et al., 1992). The microbial
biomass C was calculated as [2.64  (K2SO4-extractable Physical and chemical soil characteristics were affected by
organic C in fumigated soil  K2SO4-extractable organic C in charcoal production at the kiln site. Soil pH showed this trend:
unfumigated soil)] (Vance et al., 1987) and the microbial kiln > transition zone > control (Table 1). The organic C and
biomass N as [2.22  (K2SO4-extractable total N in fumigated total N contents of soil were lower in the TRANSITION ZONE
soil  K2SO4-extractable total N in unfumigated soil)] (Jenkin- and KILN treatments than in the CONTROL treatment
son, 1988). (P < 0.05).
The Ca2+ content was intermediate in the TRANSITION
2.5. Aerobic incubation ZONE sites compared to the CONTROL and KILN treatments
(Table 2). Extractable P showed this trend: control > transition
Twelve sub-samples of 25 g soil from each sampling location zone > kiln. The K+ content was 1.5 times higher in the KILN
were added to 120 ml glass flasks. Three were chosen at treatment compared to the TRANSITION ZONE and CONTROL
random from each treatment and soil was extracted for treatment (P < 0.05). The Mg2+ content was not affected by
inorganic N with 100 ml 0.5 M potassium sulphate (K2SO4). The treatment.
samples were shaken for 60 min and filtered through What-
man No 42 paper1 and stored at 20 8C before analysis. This 3.2. Microbial biomass C and N
provided zero-time samples. The remaining glass flasks were
placed in 945 ml glass jars containing one vessel with 10 ml The microbial biomass C was lower in the KILN treatment
distilled H2O and one beaker with 20 ml 1 M NaOH to trap CO2 compared to the TRANSITION ZONE and CONTROL treatments
evolved. The jars were sealed and stored in the dark for 28 days in the rainy and dry season (P < 0.05) (Table 3). The microbial
at 22  2 8C. An additional nine jars with one beaker of 10 ml biomass N was lower in the KILN treatment compared to the
distilled H2O and one beaker of 20 ml 1 M NaOH were sealed TRANSITION ZONE treatment in the rainy and dry season
and served as controls to account for the CO2 trapped from the (P < 0.05). The microbial biomass N was lower in the
air. After 7, 14 and 28 days, three jars selected randomly from TRANSITION ZONE treatment compared to the CONTROL
each treatment were opened and the vessel containing NaOH treatment in the rainy and dry season (P < 0.05). Although
was removed. A 5 ml NaOH aliquot was analyzed for CO2 similar trends were found in the microbial biomass for the
trapped. The soil was removed from three flasks and extracted rainy and dry seasons, there were greater differences in
for inorganic N as described for zero-time samples. All microbial biomass between the transition zone and control
remaining flasks were opened, aired for 10 min to avoid sites during the dry season.
anaerobicity, sealed and stored in the dark.
3.3. Ammonifiers, nitrifiers and denitrifiers
2.6. Statistical analyses
The number of ammonifiers was lower in the KILN treatment
Concentration of NH4+, NO2, NO3, and microbial biomass C than in the CONTROL treatment in the rainy and dry season
and N were subjected to one-way analysis of variance using (P < 0.05) (Table 4). The amount of NH4+ and NO2 oxidizers

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 Microbial biomass N
1 1
Rainy (mg kg soil) Dry (mg kg soil) Rainy (mg kg1 soil) Dry (mg kg1 soil)

KILN 2177 (44)a Bb 406 (50) B 220 (10) C 52 (6) C


TRANSITION ZONE 2969 (77) A 800 (54) A 476 (17) B 125 (10) B
CONTROL 2724 (90) A 990 (77) A 591 (35) A 213 (16) 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).
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 when soil was sampled in the dry season (P < 0.05) (Fig. 1b). The
in the CONTROL treatment in both the rainy and dry season concentration of NO2 remained <0.3 mg N kg1 soil and was
(P < 0.05). The amount of denitrifiers was lower in the KILN not affected by treatment or season (Fig. 1c). The concentration
treatment than in the TRANSITION ZONE and CONTROL of NO3 did not change over time when soil was sampled in
treatments in both the rainy and dry season (P < 0.05). the dry season, but did increase when sampled in the rainy
season except in the CONTROL treatment (P < 0.05).
3.4. Dynamics of C and N After 28 days, the amount of soil organic matter miner-
alized to N, i.e. the amount of inorganic N (NH4+ + NO2 +
The emission of CO2 was linear in the soil sampled in the rainy NO3) at the end of the incubation minus the inorganic N at the
season, but slowed down after 7 or 14 days when sampled in the onset of the incubation, was 0.28% in soil at the KILN
dry season (Fig. 1a). The lowest C mineralization was found in treatment, 0.18% in the TRANSITION ZONE and 0.10% in the
the KILN treatment. The concentration of NH4+ remained CONTROL soil in the dry season and 0.18%, 0.13% and 0.01% in
constant in soil sampled in the rainy season, but increased 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 55

4. Discussion respectively. The microbial biomass N was 4.3% of soil organic


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