Biomass and Bioenergy 83 (2015) 123e130

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Biomass and Bioenergy

journal homepage: http://www.elsevier.com/locate/biombioe

Research paper

Soil CO2eC flux and carbon storage in the dry tropics: Impact of land-
use change involving bioenergy crop plantation
Mahesh Kumar Singh a, Hastings Astley b, Pete Smith b, Nandita Ghoshal a, *
a
Centre of Advanced Study in Botany, Department of Botany, Banaras Hindu University, Varanasi 221005, India
b
Institute of Biological and Environmental Sciences, School of Biological Sciences, University of Aberdeen, AB24 3UU, Scotland, UK

a r t i c l e i n f o a b s t r a c t

Article history: The study was conducted to evaluate the impact of land-use change in the dry tropics on soil CO2eC flux
Received 8 September 2014 and soil organic carbon (SOC) storage, along with the major factors controlling them i.e. soil microbial
Received in revised form biomass (SMB), belowground net productivity (BNP), and soil aggregate size fractions. Land-use change,
31 August 2015
in this study, involved conversion of natural forest, to degraded forest and then to an agroecosystem or a
Accepted 18 September 2015
Available online xxx
bioenergy crop plantation of Jatropha curcas. The soil CO2eC flux was highest in the agroecosystem
followed in decreasing order by degraded forest, bioenergy crop plantation and smallest in the natural
forest. The inverse trend was found in case of SOC storage, SMB and BNP. The proportion of macroag-
Keywords:
Land-use change
gregate in the soil follows the trend of SOC storage, whereas mesoaggregate follows the trend of CO2eC
Climate change flux across all land-use types. The CO2eC flux showed significant negative correlation with BNP, SMB,
Carbon storage macroaggregate size fraction, and SOC. Our study suggests that the flux of soil CO2eC was regulated
Soil CO2eC flux directly by the soil aggregate fraction and not by SMB or BNP. However, soil aggregate formation was, in
Dry tropics turn, related to the SMB and/or below BNP. Macroaggregates appear to protect the SOC, which results in
Soil aggregates lower CO2eC flux and higher SOC storage. It can be concluded that the bioenergy crops plantation on
degraded forest lands in the dry tropics, may increase C storage in soil and reduce soil CO2eC flux,
thereby helping in the mitigation of global climate change in addition to providing feed stocks for fossil
fuel substitution.
© 2015 Published by Elsevier Ltd.

1. Introduction
Land-use change can lead to changes in the level of atmospheric
greenhouse gases, especially CO2, by modifying their emission from
the terrestrial ecosystems [1e3]. Currently, the contribution of
land-use change to emissions of C to the atmosphere is 1.4 Pg C yr
1
globally; which is one fifth that due to fossil fuel emissions [4].
Anthropogenic activities have overridden the natural variability in
C storage and flux [5], leading to global efforts to reduce atmo-
spheric CO2 concentrations, either by reducing anthropogenic
emissions or removing atmospheric CO2 through sequestration in
the biosphere [6,7]. Carbon storage in the soil represents the bal-
ance between accumulation and loss of carbon from soil.
Soil CO2 flux or soil respiration in any terrestrial ecosystem is the
sum of autotrophic (root) and heterotrophic (soil microbes) respi-
ration [8]. Heterotrophic respiration exceeding the carbon input
* Corresponding author.
E-mail address: n_ghoshal@yahoo.co.in (N. Ghoshal).
into the soil results in reduction in SOC. Autotrophic and hetero-
trophic soil respiration are very difficult to measure separately, but
estimates of composite soil CO2 may serve as the indicator of C
output from soil. Soil microbes, play an important role in regulating
dynamics of soil organic matter and can be represented by soil
microbial biomass, which comprises the soil microorganisms
smaller than 5  10
15 m3 [9]. A substantial amount of carbon is
allocated to the root biomass therefore belowground net produc-
tivity may give an indication of the extent of root respiration [10,11].
A major land-use change is the conversion of natural forests into
degraded forest due to excessive extraction of their products
including wood and/or non-wood products, grazing and poor
management practices [12]. Degraded forest is generally described
as the forest that has been impacted by human activity to the extent
that it no longer provides the functions of non-degraded forest.
Another major land-use change pattern is the conversion of forests
to agroecosystems which lead to alteration in the input/output
balance of SOC. Agroecosystems are anthropogenically perturbed
ecosystems and used predominantly for food or fibre production.
Among various measures for the rehabilitation of degraded

http://dx.doi.org/10.1016/j.biombioe.2015.09.009
0961-9534/© 2015 Published by Elsevier Ltd.
124 M.K. Singh et al. / Biomass and Bioenergy 83 (2015) 123e130
lands, the plantation of perennial bioenergy crop like Jatropha
curcas has been recommended [13,14], however, some conflicting
reports are also available [15]. Although many studies have covered
the impact on various soil characteristics and the biodiesel poten-
tial of J. curcas plantations worldwide [14,16e19] an extensive study
of soil C dynamics, especially carbon storage in soil and soil CO2eC
flux [20] is limited particularly in the dry tropics. The evaluation of
C storage in soil and seasonal variations in soil CO2eC flux due to
land-use changes in the dry tropics is required to better understand
carbon dynamics in this climate.
Soil is referred to as a ‘sink’ when it stores C and a ‘source’ when
it loses C [21], and changes in the role of soil from ‘source’ to ‘sink’
or vice-versa depends mainly on the land-use, which defines the
annual organic material input among various other factors. Any
land-use management strategy to enable the soil act as ‘sink’ may
eventually help in mitigation of global climate change. The
assessment of soil CO2 flux has attracted interest worldwide as it is
a major factor of carbon dynamics in the terrestrial ecosystem, and
can be taken as the indicator of carbon availability in soil for
decomposition [22], and the transformation of soil organic carbon
to atmospheric CO2 [23]. Many studies evaluated the effect of cli-
matic factors on soil CO2 flux [1,24e27] biotic and abiotic factors
[28], but a study on the impact of land-use changes on soil CO2eC
flux is lacking, particularly with reference to the role of soil struc-
ture in the dry tropics.
Soil structure plays an important role in the storage of carbon.
Degradation of soil due to land-use change is reported to be
strongly related to changes in soil structure [29]. Soil structure is
often represented through the aggregate size fraction of soil. Soil
aggregates are the group of soil particles bound with each other by
adhesive force more strongly than surrounding particles, which is
formed by a constructive process starting from primary particles to
microaggregate (<0.250 mm), mesoaggregate (0.250e1.00 mm)
and macroaggregate (>1.00 mm) [30]. Soil aggregates protect the
soil organic matter from decomposition by trapping and mini-
mizing the direct contact of soil organic matter to soil microor-
ganisms [30], and provide stability to soil structure. Distribution of
aggregate size fraction in dry soil, therefore, can be used as an index
of soil disturbance and is helpful to identify sustainable soil man-
agement [31].
In the current study, we hypothesized that (i) land-use change
can influence the rate of soil CO2eC flux and alters the soil status of
carbon storage, (ii) the rate of soil CO2eC flux and carbon storage
depends not only on the soil microbial biomass and belowground
net productivity, but also on the soil structure in terms of soil
aggregate size fraction, and (iii) the planting of bioenergy crops like
J. curcas on degraded forest lands in the dry tropics might increase
the soil carbon stock through enhanced storage of soil organic
carbon. To test these hypotheses, this study aimed at evaluating the
impact of land-use change from natural forest to degraded forest
and then to either bioenergy crop plantation, or agroecosystems in
the dry tropics in terms of (i) seasonal variations in soil CO2eC flux,
(ii) soil carbon storage, (iii) variations in soil microbial biomass, (iv)
belowground net productivity, (v) changes in the proportion of soil
aggregate size fractions, and (vi) the factors controlling soil CO2eC
flux.
2. Material and methods
2.1. Study site
The four adjacent land-use types selected for study were: nat-
ural forest (10 km2), degraded forest (4.5 km2), agroecosystem
(0.0625 km2) and bioenergy crop plantation in the form of J. curcas
plantation (0.16 km2). The degraded forest, agroecosystem and
bioenergy crop plantation which had initially been natural forest,
were situated at Rajeev Gandhi South Campus, Banaras Hindu
University, Barakachha Mirzapur, Uttar Pradesh, India, while the
natural forest was in the Marihan range, about 7e8 km away from
Barakachha Mirzapur. The study was conducted during September
2011 to June 2012. The sites are at 25.1500 N latitude, 82.5800 E
longitude, and about 81 m above mean sea level. The climate of the
study region is dry tropical monsoonic with marked seasonality.
The year can be divided into three distinct seasons: rainy (July to
September), winter (November to February), and summer (April to
June); October and March are the transitional months. The annual
average rainfall was about 1876 mm (1700e2052 mm), 95% of
which fall during the rainy season. Mean monthly minimum and
maximum temperature ranged from 14.2 to 32.5  C and 25.5 to
42.8  C, respectively. The reddish to reddish-brown soil of the re-
gion was categorized as residual ultisol in the United States
Department of Agriculture soil taxonomy, sandy to sandy loam in
texture and was common to each site. The soil parent material is
derived from the Kaimur sandstones, i.e. Dhandraulorthoquartzite,
for all the land-use types. The basic soil properties of all the sites are
listed in Table 1. As can be seen from the table Ultisols are mildly
acidic.
The forest site represented the mixed dry deciduous type
dominated by Acacia catechu Wild., Albizia odoratissima Benth.,
Acacia nilotica (L.) Willd., Boswellia serrata Roxb., Nyctanthes
arbortristis L., with scattered trees of Azadirachta indica Juss. and
Zizyphus glaberrima Santap. The forest floor was covered with
herbaceous vegetation of Ocimum americanum L., Pisum arvense L.,
Rhynchosia minima (L.) DC., Cassia sophera (L.) Roxb., Acrocephalus
indicus (Burm. f.) Kuntze.,Cynodon dactylon L., Oplismenus bur-
mannii Ritz. The degraded forest site was dominated by Zizyphus
glaberrima Santap, Chrysopogan fulvus Spreng., Heteropogan con-
tortus L., Adina cordifolia Roxb. and scattered trees of Butea mono-
sperma Lamk. Herbaceous vegetation in the degraded forest was
dominated by Cassia tora L., Oldenlandia diffusa Roxb., Sporobolus
spp. Panicum psilopodium Trin., Alysicarpus varginalis (L.) DC. The
cultivation of degraded forest started in 1979 and was maintained
until the present day at the Rajeev Gandhi South Campus, Bar-
akachha. The crop sequence studied was Oryza sativa (var. HUR
3022)ebarley (Hordeum vulgare var. Manjula)esummer fallow.
Since June 1990, chemical fertilizer in the form of urea, single super
phosphate and muriate of potash at the rate of 100 N, 60 P and
40 kg K ha
1 respectively, have been applied annually. For the
rehabilitation of degraded land resulting from degradation of nat-
ural forest, the establishment of J. curcas plantations began about
ten years ago in a 100 ha area at Rajeev Gandhi South Campus. The
major objective of the plantation of J. curcas was to control soil
degradation in addition to biofuel production. J. curcus was planted
in rows with an inter row distance of 2 m and the distance between
the plants in a row was also 2 m. Evolvulus nummularis L., Glinus
oppositifolius (L.) DC., Tephrosia purpurea L., and C. tora L. were the
major herbaceous vegetation among J. curcas plantation.
2.2. Measurements
Each land-use type site was first divided into three study sites,
which were further sub-divided into nine sub-sites for natural
forest, degraded forest and bioenergy crop plantation and eight for
the agroecosystem; out of which, five were randomly selected for
sampling together at the beginning of experiment. Details about
land-use types, study sites and sub-sites are given by Singh and
Ghoshal, 2014 [32]. For estimation of soil aggregate size fraction
distribution, microbial biomass and soil organic carbon, two soil
samples were collected using the soil corer with a diameter of 4 cm
and depth of 10 cm from each sub-site (i.e., total of 10 soil samples
 M.K. Singh et al. / Biomass and Bioenergy 83 (2015) 123e130
Table 1
Variation in soil properties of the various land use types i.e. NF (natural forest), DF (degraded forest), AG (agroecosystem) and JP (Jatropha plantation). Values are mean ± SE.
LSD values are at p < 0.05.
Soil parameters Land use types
NF DF
SOC (%) 0.87 ± 0.04 0.47
TN (%) 0.140 ± 0.002 0.069
BD (g cm
3) 1.09 ± 0.03 1.21
WHC (%) 47.41 ± 2.83 43.68
pH 6.38 ± 0.07 6.78
Code: SOC ¼ Soil organic C, TN ¼ Total soil N, BD ¼ Bulk density, WHC ¼ Water holding capacity.
from each study site), and mixed together to represent the single
composite sample for a study site. From the same sub-sites, sam-
pling was done for estimation of bulk density, root biomass and soil
CO2eC flux, and the values averaged to represent a study site.
Dry aggregate size distribution was determined by dry sieving
[33]. Soil was sampled from sub sites with the help of soil corer
(4 cm dia. and 10 cm height) only in the summer season. Soil field-
moist samples were spread on blotting paper to a height of nearly
1 cm, and allowed to air dry for one week. Soil samples (100 g) were
placed on top of the nest of sieves and sieved for 3 min on an end-
to-end shaker (92 rpm), and three dry aggregate size classes
separated were,>1000 mm (macroaggregate), 212e1000 mm (mes-
oaggregate) and <53e212 (microaggregate).
Soil organic carbon concentration was analysed from the soil
sampled for dry aggregate analysis, after sieving through a 2 mm
mesh, by using dichromate oxidation and back titration of unused
dichromate [34]. The results are expressed on a dry weight basis.
Soil bulk density was estimated by removing the known volume of
soil with the help of soil corer (3 cm diameter and 10 cm height),
and dried for 24 h at 105  C.
Soil carbon storage was calculated by using the equation,
Cs ¼ SOC  BD  D  (1
RF) [35], wherein Cs is the soil organic
carbon storage (Mg ha
1); SOC in %; BD the bulk density (g cm
3); D
the soil depth (cm), and RF the volumetric fraction of rock frag-
ments which are >2 mm (%). In the soil samples of the present
study, there were no gravel fractions, so we ignored RF factor, and
Cs was calculated by using the following equation:
Cs ¼ SOC  BD  D.
Surface soil CO2eC flux was determined during rainy, winter
and summer seasons by alkali absorption method [36] by inserting
the PVC (Poly Vinyl Chloride) chambers (height 23 cm and diameter
14 cm) in the soil up to 3 cm depth, and left overnight to stabilize. A
100 ml of beaker having 25 ml NaOH of 0.5 N with few drops of
phenolphthalein was kept on a metal stand above the soil surface in
each cylinder. Each cylinder was tightly sealed and closed on top
with cling film. The sets were left undisturbed for 24 h in the field.
The CO2eC flux was estimated by titrating excess of NaOH with HCl
after adding BaCl2 immediately after sampling. CO2eC was esti-
mated using conversion formula CO2eC]CO2/3.67.
Soil samples, in addition to those collected for soil texture, SOC
and BD were collected during rainy, winter and summer season by
using a soil corer (3 cm diameter and 10 cm height) and sieved
through a 2-mm mesh screen after removing the visible plant
debris. Soil microbial biomass C (MBC) was analysed by chloroform
fumigation-extraction method using purified CHCl3 treatment [37].
The dichromate digestion method was adopted for the determi-
nation of soil microbial biomass C and calculated as:
MBC ¼ 2.64  EC, where EC represents the difference between
organic C extracted in the K2SO4 extracts of fumigated and non-
fumigated soils. The microbial biomass C was analysed during
rainy, winter and summer season.
For estimation of root biomass, soil monoliths were excavated
125
LSD
AG JP
± 0.01 0.29 ± 0.03 0.71 ± 0.02 0.08
± 0.004 0.064 ± 0.003 0.083 ± 0.002 0.01
± 0.02 1.31 ± 0.03 1.16 ± 0.03 0.08
± 3.06 39.10 ± 1.48 45.39 ± 2.83 8.56
± 0.06 7.02 ± 0.10 6.64 ± 0.15 0.33
with the help of soil corer (10 cm dia. and 10 cm height) during
rainy, winter and summer seasons. For the retrieval of roots, soil
monoliths were then washed with the help of a fine water jet over a
twin sieve set (2 mm mesh top and 0.5 mm mesh below). The
recovered root mass were air dried, cleaned and oven dried at 80  C.
The sum of successive increment of plant root biomass with season
is considered as belowground net productivity. The sampling dates
were 14, 15 September, 20, 21 January and 14, 15 June for rainy,
winter and summer season, respectively.
2.3. Statistical analysis
SPSS package (version 16), R and Minitab (V 16) were used for
data analysis. The statistical unit was study site with N ¼ 3 for each
parameter during each season of a site. The effect of land-use type
and season on the rate of soil CO2eC flux was tested using two-way
ANOVA. The Duncan test was performed to observe the significant
difference. Pearson's correlation test was performed for the corre-
lation of annual mean of soil CO2eC flux and soil carbon storage
with different parameters. All values are expressed as mean ± SE.
Means of data were compared using least square difference (LSD) at
p < 0.05, p < 0.01 and p < 0.001.
The impact of land use on the soil macro, meso and micro-
aggregate fractions was tested using a one way ANOVA.
Considering soil CO2eC flux as the predicted variable and soil
aggregate mass fractions and associated SOC, total SOC, below-
ground net productivity, pH and soil microbial biomass as covariate
explanatory variables and season and land-use as factors, a data
exploration eliminated collinear variables and a stepwise GLM was
performed.
Finally using the cohort soil model [38,39] the transition of
degraded land to J. curcas plantation was modelled, and the annual
change in organic carbon per year estimated. This was achieved by
using the degraded land SOC as the initial value for the J. curcas
plantations SOC and matching the measured value in this experi-
ment after 10 years of Jatropha. Using plant material degradation
rates for woody biomass (98% input with a decomposition time
constant of 15 years and 2% with 500 years). The history match was
achieved by varying the annual organic material input into the soil.
The model was used to estimate the SOC after 50 years.
3. Results
3.1. CO2eC flux
The flux of soil CO2eC (82e171 mg CO2eC m
2 h
1) was the
highest in agroecosystems, and decreased substantially in degraded
forest, bioenergy crop plantation, and attained its minimum in the
natural forest for all the seasons (Fig 1a). The variation in soil CO2eC
flux throughout the season was not significant, though a trend was
evident for maximum during the rainy season, and decreased
gradually through winter to attain its minimum during summer in
126 M.K. Singh et al. / Biomass and Bioenergy 83 (2015) 123e130

Fig. 1. (a) Variation of soil CO2eC flux (mean ± SE) in four land-use types i.e. natural
forest (NF), degraded forest (DF), agroecosystem (AG) and Jatropha plantation (JP)
during rainy, winter and summer seasons. Annual mean of soil CO2eC flux of the three
seasons of each land-use type was indicated by horizontal line and (b) seasonal vari-
ation of soil microbial biomass C (mean ± SE) of four land use types during the three
season. Different alphabetic lower case letters represent the significant difference
among the land use types (Duncan test, N ¼ 3, p < 0.05); whereas different numerical
represents the significant difference among the season (Duncan test, N ¼ 3, p < 0.05) in
both the figures.

the natural forest, degraded forest or bioenergy crop plantation.
However, the agroecosystems showed an altogether different trend
with the maximum flux during rainy season that decreased during
winter but slightly increased during summer. It is evident from the
analysis result of the two-way ANOVA that both land-use type
(f ¼ 38.96, df ¼ 3, p < 0.001) and season (f ¼ 9.66, df ¼ 2, p < 0.001),
influenced soil CO2eC flux, more than the interaction between
land-use type and season (f ¼ 0.80, df ¼ 6, p > 0.05).
3.2. Aggregate size fraction
Soil aggregate size fractions varied distinctly with land-use type
as evident form a one way ANOVA. The proportion of macroag-
gregates varied with land use (f ¼ 5.7, df ¼ 3, p < 0.022) and was
highest in soils of natural forests (69%) and was successively lower
in bioenergy crop plantation (62%), degraded forest (59%) and
agroecosystems (51%). The proportion of mesoaggregate showed
the opposite trend (f ¼ 12.85, df ¼ 3, p < 0.002), the proportion was
largest in the agroecosystem (36%) followed by a decreasing order
for degraded forest (27%), bioenergy crop plantation (21%) with the
minimum in natural forest (18%). There was no significant differ-
ence between land uses for microaggregates (f ¼ 0.77, df ¼ 3,
p < 0.541). The impact of land-use change on aggregate size fraction
is shown graphically in Fig 2c.
3.3. Soil carbon storage
Soil carbon storage significantly differed with land-use type and
ranged from 3.78 to 9.47 Mg ha
1. It attained its maximum in the
natural forest, followed by a decreasing trend for bioenergy crop
plantation, degraded forest with the minimum in agroecosystems
(Fig. 2b). Although SOC was measured on the mixed total sample, if
the SOC value was applied to the proportions of the aggregate, the
Fig. 2. (a) Belowground net productivity (mean ± SE) of the four land use types i.e.
natural forest (NF), degraded forest (DF), agroecosystem (AG) and Jatropha plantation
(JP) at the upper layer (0e10 cm) of soil; (b) soil C storage (mean ± SE) of the four land
use types at the upper layer of soil (0e10 cm) of soil and (c) proportion of aggregate
size fractions (mean ± SE) in four land-use types. Different alphabetic lower case
letters represent the significant difference among the land use types (Duncan test,
N ¼ 3, p < 0.05) in each figure.
SOC in the macroaggregate and microaggregate varied with land-
use, but the mesoaggregate SOC was constant.
3.4. Soil microbial biomass C
Soil microbial biomass C ranged from 257 to 723 mg
1 with its
maximum level in natural forest, and decreased through bioenergy
crop plantation, degraded forest with the minimum in the agro-
ecosystem. Highest level of soil microbial biomass was recorded
during summer and lowest during the rainy season across all land-
use types (Fig. 1b).
3.5. Belowground net productivity
Belowground net productivity varied significantly with land-use
type and ranged from 783 to 3253 kg ha
1. Maximum belowground
net productivity was found under natural forest followed by a
decreasing trend for bioenergy crop plantation, degraded forest
and agroecosystem (Fig. 2a).
The CO2eC flux showed strong negative correlation with soil
carbon storage (r2 ¼ 0.83, p < 0.01, n ¼ 12) and soil microbial
biomass C (r2 ¼ 0.70, p < 0.01, n ¼ 12) (Fig. 3). The annual mean
of CO2eC flux showed a negative relationship with belowground
net productivity (r2 ¼ 0.925, n ¼ 4, p < 0.05), soil macroaggre-
gate (r2 ¼ 0.942, n ¼ 4, p < 0.05) and a positive relationship with
mesoaggregate size fraction (r2 ¼ 0.973, n ¼ 4, p < 0.05),
 M.K. Singh et al. / Biomass and Bioenergy 83 (2015) 123e130 127

Fig. 3. (a) Relationship of soil CO2eC flux and soil microbial biomass C across the four land use types and three seasons; (b) relationship of soil C storage with microbial biomass C
across the four land use types and three seasons and (c) relationship of soil CO2eC flux and soil C storage across the four land use types and three seasons.

whereas microaggregates showed no relationship (r2 ¼ 0.008,

n ¼ 4, p > 0.05) with soil CO2eC flux across all the land-use
types.
Soil carbon storage across all the land-use types through the
three seasons showed a strong negative correlation with the soil
CO2eC flux (r2 ¼ 0.83, p < 0.01, n ¼ 12) (Fig. 3), but a weak negative
relationship with soil microbial biomass C (r2 ¼ 0.40, p > 0.05,
n ¼ 12) (Fig. 3).
The step wise GLM showed that the CO2 flux rate depended on
both SOC and season, which improved the R-Sq (adj) to 95%
(p > 0.000). pH was not used in this model at was co-linear with
SOC, however an alternative GLM model showed that CO2 flux rate
depend on pH R-Sq (adj) to 95% (p > 0.000) and season did not
improve the model.
Annual mean of soil C storage showed a positive relationship
with macroaggregate (r2 ¼ 0.934, n ¼ 4, p < 0.05), belowground net
productivity (r2 ¼ 0.913, n ¼ 4, p < 0.05) and a negative relationship Fig. 4. Carbon co-hort model using the initial point as the degraded land and a leaf
with mesoaggregate size fraction (r2 ¼ 0.978, n ¼ 4, p < 0.05), but input of 0.66 Mg ha
1 and fitting the measured value at 10 years.
there was no relationship with microaggregate size fraction
(r2 ¼ 0.015, n ¼ 4, p > 0.05).
The Cohort model matched the CO2 emission for both the initial 4. Discussion
degraded forest and J. Curcas plantation after 10 years by estimating
an annual input of organic carbon of 0.66 Mg and the equilibrium The major contribution to soil CO2eC flux is through respiration
SOC was similar to that of a natural forest (Fig. 4). of roots and soil microbes [28] and also through the mineralization
128 M.K. Singh et al. / Biomass and Bioenergy 83 (2015) 123e130
of soil organic matter. In the present study, a strong negative
relationship was found among soil CO2 flux and soil microbial
biomass and soil carbon storage across all the land-use types and
season. Such observations are contrary to most other studies, which
reported a direct relationship among soil CO2eC flux and soil mi-
crobial biomass and root biomass [28,40,41]. The higher CO2 flux
also correlated with the higher pH. Ultisols are usually acidic but
the addition of Urea, super phosphates and muriate of potassium as
fertilizers in the agroecosystem would tend to increase pH as would
the animal droppings in the degraded forest. This may create a
priming effect for the heterotrophic rhizo-bacteria and increase the
population of which would be limited by the organic substrate. The
lower pH and higher carbon input in the natural forest, and to a
lesser extent the J. curcas plantations, would tend to protect the
organic material and create a more diverse, and thus larger,
rhizosphere ecosystem.
Although soil microbial biomass and belowground net produc-
tivity was highest in natural forest compared to other land-use
types, the least soil CO2eC flux was observed in the forests in this
study (Fig. 1a). In the natural forest, a higher proportion of soil
macroaggregate compared to mesoaggregate (Fig. 2c) is attributed
to the presence of the higher soil microbial biomass and below-
ground net productivity. Exudates of root and soil microbes act as a
gluing agent for aggregation of soil particles and thus facilitate the
formation of macroaggregates [42,43], which in turn protects
organic matter from mineralization via the formation of organo-
emetallic complexes [29]. In spite of the maximum level of soil
microbial biomass in natural forest, the lowest contribution to total
soil CO2eC flux might be due to the presence of more stable, less
dynamic microbial biomass, which was reported to occur in eco-
systems having less stress and more favourable conditions [44,45].
The presence of a higher proportion of macroaggregates in this
forest, might also protect soil organic matter within the aggregates
and prevent mineralization, resulting on one hand in lower soil
CO2eC flux, and on the other in higher soil C storage [46]. In
addition, the priming effect on microbial activity of fertilizer
application was not available in the natural forest or J. curcas
plantations, whist in the agroecosystem, due to moderate chemical
fertilizer application is was, and to a lesser extent in the degraded
forest, due to animal droppings.
The major drivers of degradation of natural forests in the dry
tropics are the logging of trees for various purposes and cattle
grazing. In the present study, a considerable decrease (71%) in
belowground net productivity was observed in degraded forest
compared to natural forest, attributed to a severe loss of vegetation
cover (Fig. 2a). Reductions in the organic inputs to soil in the form
of above- and below-ground biomass resulted in the significant
decrease in soil microbial biomass compared to natural forest in
this study [32]. In spite of considerable reductions in soil microbial
biomass and belowground net productivity, a considerable increase
in soil CO2eC flux was observed (Fig. 1a). The regular disturbance
might have led to microbial biomass becoming less stable and more
dynamic, which probably resulted in higher microbial respiration
rate in terms of higher soil CO2 flux [44]. In the present study on the
degraded forest, the proportion of mesoaggregates increased (52%)
compared to natural forest, and that of macroaggregates decreased
by 14% (Fig. 2c), probably due to degeneration of soil macroaggre-
gates to form meso- and microaggregates mainly through tram-
pling by cattle [47]. In addition, lower microbial biomass and also
low belowground net productivity perhaps provided less favour-
able conditions for the formation of macroaggregates. The lower
proportion of macroaggregates, and higher proportion of mesoag-
gregates compared to natural forest perhaps facilitated minerali-
zation of protected soil organic matter resulting in higher soil CO2
flux and lower C storage in soil.
Tillage practices disrupt the soil aggregate into smaller fractions
and thus facilitate formation of meso- and microaggregates [48].
The major fraction of the aboveground biomass is periodically
harvested from the agroecosystem, leaving behind the small input
of organic matter through crop roots and stubble, which might
explain the lower level of soil microbial biomass in the agro-
ecosystem [32]. Root exudates that help in formation of macroag-
gregates are produced only from live roots. The life of which is of
limited duration in the agroecosystem as rapid root mortality oc-
curs following the grain forming stage [49], as a result it is also
expected that root exudation might not be high enough to glue the
mesoaggregates to form macroaggregates. The limited root exu-
dates for shorter durations and the destruction of macroaggregates
through ploughing might be the reason for higher proportion of
mesoaggregates to macroaggregates in the agroecosystem (70%)
compared to the natural forest (26%) (Fig. 2c). The higher propor-
tion of mesoaggregates compared to macroaggregates lead to the
exposure of soil organic matter to microbes which, in turn, in-
creases the probability of mineralization of soil organic matter [50],
which could also be the reason of higher soil CO2 flux and lower
storage of C in soil. Elliott [51] reported more soil CO2eC efflux from
crushed macroaggregates over intact macroaggregates under lab-
oratory conditions. Islam & Weil and Kristensen et al. [52,53] re-
ported higher rates of decomposition of organic matter and crop
residue in agroecosystems owing to a highly oxidative soil envi-
ronment compared to other land-use types, and this might also be
the reason for reduced soil C storage. It is also apparent that the
lowest soil microbial biomass might be highly dynamic and least
stable due to the stress of periodic disturbance, thereby producing
the highest soil CO2eC flux.
In the plantations of J. curcas, cattle grazing were almost negli-
gible as the leaves are not palatable. J. curcas plantations have an
extensive root system with a dense net of horizontal lateral roots
[54], which probably favours formation of macroaggregates [29].
Ogunwole et al. [14] reported an increase in the proportion of
macroaggregates by the cultivation of J. Curcas on degraded soil.
Almost all the nutrient-rich leaves shed during winters were added
to the soil as leaf litter, and thus helped to maintain higher soil
microbial biomass [32]. Lower levels of disturbance, due to limited
cattle grazing and more organic input to the soil through leaf litter
compared to degraded forest or agroecosystems could have facili-
tated the occurrence of more stable, less dynamic microbial
biomass, which could explain the lower soil CO2 flux. Higher root
biomass and microbial biomass along with a greater proportion of
macroaggregates might have created conditions conducive for high
storage of C in soil and so also for less efflux of CO2eC compared to
the degraded forests and agroecosystems. The study suggests that
the plantation of bioenergy crops stored more C and permitted less
soil CO2eC flux compared to degraded forests or agroecosystems.
It is evident that the flux of soil CO2eC was regulated directly by
the soil aggregate fraction and not by the soil microbial biomass or
belowground net productivity. However, soil aggregate formation
was in turn, also regulated by the soil microbial biomass and/or
belowground net productivity. A positive feedback loop among the
soil microbial biomass, belowground net productivity and macro-
aggregates might control the soil CO2eC flux and C storage in soil,
instead of any one factor in isolation.
The general conception is that the flux of soil CO2eC is regulated
more by the biotic variables i.e., soil microbial biomass and plant
biomass compared to soil physical and chemical properties [28,55].
However, in the present study, the role of macro- and meso-
aggregate were no less important than that of the biotic variables,
as shown by the relationship of soil CO2eC flux and soil microbial
biomass, belowground net productivity and aggregate size fractions
of soil. Moreover, in most previous studies, the biotic variables were
 M.K. Singh et al. / Biomass and Bioenergy 83 (2015) 123e130
positively correlated with soil CO2eC flux [28], whereas in the
present study, soil CO2eC flux was inversely proportional to not
only the soil macroaggregate size fraction but also the soil microbial
biomass, belowground net productivity, and directly related to the
mesoaggregate fraction of soil, whose formation was enabled by
the lack of priming effect from pH increasing input and disturbance
by tillage.
Although soil CO2eC flux did not show significant variation
through seasons, a definite trend was evident in natural forest,
degraded forest and bioenergy crop plantation (Fig. 1a), and this
could be attributed to seasonal variations in root respiration rather
than the soil microbial biomass, as the trend of soil CO2eC flux was
similar to that of former. Belowground net productivity based on
the root biomass was maximum during rainy season and minimum
during summer, indicating maximum belowground net produc-
tivity during the former. The soil microbial biomass was reported to
be maximum during summer and minimum during the rainy sea-
son [32]. In the agroecosystem, live roots remained for shorter
duration in the soil during rains as rice roots, and in winter as barley
roots. Rice had higher root biomass compared to barley. As the rice
crop was harvested the dead root mass was left in the soil, to
decompose slowly until the next rainy season [56]. It is evident that
after the barley harvest, i.e. at the onset of summer, dead root mass
was added to the soil. It was during the summer that decomposing
roots and barley roots together could have contributed to the in-
crease in soil CO2eC flux. Sugihara et al. [26] also reported low CO2
flux during rainy season relative to dry season in the
agroecosystem.
Since soils may act both as ‘source’ and ‘sink’ of carbon, any
variations due to changes in land-use pattern may alter their course
and this in turn, may also have an impact on the climate change.
This study shows that when natural forests are degraded there are
significant losses of carbon in terms of decrease in soil organic
carbon stock, and an increase in soil CO2eC flux, which perhaps
resulted in the soils acting as a source. When degraded forests
converted to agroecosystems, the soil organic carbon was lower,
with a large increase in soil CO2eC flux, suggesting that this con-
version will be a major carbon ‘source’. Plantation of bioenergy
crops on degraded forest land increased soil organic carbon stocks
significantly, and decreased the soil CO2eC flux considerably within
ten years of plantation, resulting the soils acting as a carbon ‘sink’.
On the basis of the soil carbon model, using the initial point of
soil carbon as that of degraded land and a leaf input of 0.66 Mg ha
1,
fitting the measured value at 10 years, indicated that the equilib-
rium SOC after 50 years would be around 12 Mg ha
1which is
similar to that of natural forest (Fig. 4). The cohort model is
computationally similar to the RothC model and requires knowl-
edge of the organic matter input and input degradation rate,
neither of which were measured in this experiment. The input was
estimated by fitting to the observed SOC levels and use of generic
decomposition time constants for woody biomass. To improve the
model, future work is needed to determine actual decomposition
rates for these environmental conditions. However, notwith-
standing these caveats it is estimated that the J. curcas plantation
may store as much soil carbon as a forest, as well as providing a
feedstock which will offset fossil fuel use.
5. Conclusion
In conclusion, the plantation of bioenergy crops such as
J. curcas in the dry tropics may be a potential rehabilitation
strategy for degraded lands, as it can increase C storage in soils and
reduce soil CO2eC efflux, thereby helping in the mitigation of
climate change. This is in addition to the carbon mitigated by the
displaced fossil fuels emissions. Thus plantation of bioenergy
129
crops can have value in rehabilitating degraded lands not only in
terms of economic sustainability, but also in terms of environ-
mental sustainability.
Acknowledgements
We thank the Head and the Coordinator, Centre of Advanced
Study in Botany, Department of Botany, for providing laboratory
facilities. University Grants Commission, New Delhi, India provided
financial support in form of University Research Fellowship to Mr.
Mahesh Kumar Singh.
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