Soil Biology & Biochemistry 36 (2004) 2111–2114

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Short communication
Effects of root and litter exclusion on soil CO2 efflux and microbial
biomass in wet tropical forests
Yiqing Lia,*, Ming Xua, Osbert J. Sunb, Wangcheng Cuic
a
Department of Ecology, Evolution and Natural Resources, Rutgers, The State University of New Jersey,
14 College Farm Road, New Brunswick, NJ 08901-8551, USA
b
Laboratory of Quantitative Vegetation Ecology, Institute of Botany, The Chinese Academy of Sciences, Beijing 100093, China
c
Xinjiang Institute of Ecology and Geography, Chinese Academy of Sciences, 40-3 Beijing South Road, Ulmuqi, Xinjiang 830011, China
Received 30 December 2003; received in revised form 6 June 2004; accepted 15 June 2004

Abstract
We examined the effects of root and litter exclusion on the rate of soil CO2 efflux and microbial biomass at a soil depth of 25 cm in a
secondary forest (dominated by Tabebuia heterophylla) and a pine (Pinus caribaea) plantation in the Luquillo Experimental Forest in Puerto
Rico. The experimental plots were initially established in 1990, when root, forest floor mass and new litterfall were excluded for 7 y since
then. Soil respiration was significantly reduced in the litter and root exclusion plots in both the secondary forest and the pine plantation
compared with the control. Root exclusion had a greater effect on soil CO2 efflux than the litter exclusion in the plantation, whereas a reversed
pattern was observed in the secondary forest. The reduction of microbial biomass in the root exclusion plot was greater in the secondary
forest (59%) than in the plantation (31%), while there was no difference of the reduction in the litter exclusion plots between these forests.
Our results suggest that above-ground input and roots (root litter and exudates) differentially affect soil CO2 efflux under different vegetation
types.
q 2004 Elsevier Ltd. All rights reserved.

Keywords: Soil CO2 efflux; Trenching; Litter removal; Plantation; Secondary forest; Microbial biomass

Large amounts of C are released into the atmosphere as
CO2 during the decomposition of litter added to soil from
above-ground and below-ground sources. Raich and
Nadelhoffer (1989) estimated that the above-ground litter
inputs in many forest ecosystems contribute approximately
33% of the annual C loss through soil CO2 efflux, suggesting
that the above-ground litter input exerts an important
influence on soil C dynamics. Identification of the
contribution to total soil CO2 efflux by below-ground
sources, however, has proven to be considerably more
difficult. For example, root contribution to total soil CO2
efflux ranges from 30 to 93% (Nakane et al., 1983; Bowden
et al., 1993; Laudelout and Thierron, 1996; Ryan et al.,
1997; Xu et al., 2001). Bowden et al. (1993) found that the
contribution of CO2 from below-ground litter accounted for
* Corresponding author. Tel.: C1-732-932-0640; fax: C1-732-932-
8746.
E-mail address: yiqingli@crssa.rutgers.edu (Y. Li).
0038-0717/$ - see front matter q 2004 Elsevier Ltd. All rights reserved.
doi:10.1016/j.soilbio.2004.06.003
30% of the total in a temperate mixed hardwood forest. In
addition, the rates of soil CO2 efflux vary largely among
major biomes through different above- and below-ground
litter input and decomposition (Schlesinger, 1977; Raich
and Schlesinger, 1992; Raich, 2000). Therefore, the relative
importance of above- and below-ground litter input in
regulating soil CO2 efflux in different forest ecosystems is
critical to the understanding of C cycles at landscape and
larger spatial scales. In this study, we used a pine plantation
and a secondary forest of similar age class to examine the
effects of root and above-ground litter input on soil
respiration and microbial activities.
This study was conducted between 1996 and 1997 in the
Luquillo Experimental Forest in north-eastern Puerto Rico,
and consisted of parallel measurements in a pine plantation
and a secondary forest. The site receives an average annual
precipitation of 3500 mm with annual mean temperature of
22.3 8C (Brown et al., 1983). The plantation was established
on an abandoned cropland in 1976 as part of a reforestation
2112 Y. Li et al. / Soil Biology & Biochemistry 36 (2004) 2111–2114

program of the United States Forest Service (Lugo, 1992).

The secondary forest was naturally developed on the same
abandoned cropland during the same period. The plantation
is dominated by Pinus caribeae with few shrub and grass
species on forest floor. The secondary forest consists of a
sparse overstory, dense understory, abundant shrubs,
grasses, and ferns. Three blocks were arranged into a square
plot of 0.25 ha in the plantation (5.6 ha of the total area) and
in the secondary forest (7.2 ha) initially established by the
SOMD project in 1990. Three treatments (root exclusion,
litter exclusion, and root-and-litter exclusion) and a control
plot were established in each block. Within each treatment
including the control, we randomly selected four sampling
locations as independent replicates for soil CO2 efflux and
microbial biomass measurement. The litter exclusion
treatment was implemented by building a tent 3 m!3 m
(2 m above the ground surface) using mesh screen with
openings of 2!2 mm. The root exclusion treatment was
applied by trenching soil along the four sides of the plot to a
depth of 1 m and then a car tarp sheet was buried in the soil
to keep roots growing into the plots. The litter-and-root
exclusion treatment was conducted by building a tent and
burying a car tarp sheet as described above. In addition to
the initial four treatments, new litter removal treatment was
added to the experiment in July 1996 by constructing a
rectangular tent (0.5 m2 in area and 0.65 m in height) as a
subset in each of the control plots. Litter was initially
removed in the litter removal plots in July 1996 when the
tents were first established and the litter intercepted by the
Fig. 1. Mean soil CO2 efflux (g C mK2 dK1) in the control plot, 7-year litter
tents was routinely removed throughout the study to allow exclusion (L-exc), root exclusion (R-exc) and litter-and-root exclusion
sunlight to penetrate. Soil respiration was measured every 2 (L&R-exc) plots in the plantation and the secondary forest from August
months from August 1996 to June 1997 using alkali- 1996 to June 1997. The means (GSE, nZ6) were significantly (PZ0.04)
trapping techniques following the procedure of Carter different between the treatments by Scheffe’s multiple range test.
(1993). Ground litter was not removed when soil CO2
than the plantation. The rates of soil CO2 efflux were
efflux was measured in the control plots. Microbial biomass
reduced from 2.33G0.12 to 1.02G0.07 g C mK2 dK1 and
was measured using a fumigation-incubation procedure
from 2.66G0.16 to 0.81G0.06 g C mK2 dK1 in the
(Jenkinson and Powlson, 1975) in August 1996 and March
plantation and the secondary forest, respectively, and
1997, which represented a wet (August) and a dry (March)
accounted for 70 and 56% of reduction, respectively. The
season, respectively. The treatment effect was examined and
rate of soil CO2 efflux in root exclusion plots in the
analyzed with one-way ANOVA followed by Scheffe’s
multiple range test at PZ0.05 (Montgomery, 1991). plantation decreased during the dry season. In the litter-
Soil CO2 efflux decreased considerably in both the and-root exclusion plots, soil CO2 efflux was reduced by
plantation and the secondary forest without above-ground an average of 82% in the plantation and the secondary
litter input and/or root exclusion after 7 y (Fig. 1). forest. There was no seasonal pattern of soil CO2 efflux
Litter exclusion had a greater effect on soil CO2 efflux in in both the plantation and the secondary forest.
the plantation than the secondary forest with a reduction The rates of soil CO2 efflux were lower in the new litter
of 68 and 54%, respectively. The rates of soil CO2 efflux removal plots than the control plots in both the plantation
(meanGSE) was reduced from 2.33G0.12 to 0.75G and the secondary forest (Fig. 2). The rates of soil CO2
0.06 g C mK2 dK1 and from 2.66G0.16 to 1.21G efflux were reduced by 25% in the plantation, and 15% in
0.11 g C mK2 dK1 after 7-year litter exclusion in the the secondary forest. Seasonal patterns were found in both
plantation and the secondary forest, respectively. The the plantation and the secondary forest.
rates of soil CO2 efflux in litter exclusion plots in both Total soil microbial biomass in litter exclusion plots were
the plantation and the secondary forest had no clear also reduced after 7-year litter exclusion in both the
seasonal patterns as shown in the control plots (Fig. 1). plantation and the secondary forest (Fig. 3). Litter exclusion
In contrast to the litter exclusion treatment, root reduced soil microbial biomass by 69% in the plantation,
exclusion had a greater effect in the secondary forest and 67% in the secondary forest. Root exclusion plots
 Y. Li et al. / Soil Biology & Biochemistry 36 (2004) 2111–2114
Fig. 2. Mean soil CO2 efflux (g C mK2 dK1) in the control plot and 1-year
litter exclusion (L-removal) plots in the plantation and the secondary forest
from August 1996 to June 1997. The means (GSE, nZ6) were significantly
(P!0.01) different between the treatments by Scheffe’s multiple range test.
reduced total soil microbial biomass (meanGSE) from
711G43 to 489G22 mg C kgK1 soil in the plantation, and
from 618G45 to 256G28 mg C kgK1 soil in the secondary
forest, accounting for 31 and 59% of the reduction,
respectively. In the litter-and-root exclusion plots, total
soil microbial biomass was lower than the control and other
treatments in both the plantation and the secondary forest.
Our finding of the greater percentage of reduction in soil
CO2 efflux in the plantation than in the secondary forest
after 7-year litter exclusion suggests that the responses of
soil CO2 efflux to above-ground litter exclusion vary with
vegetation types, in agreement with studies of Schlesinger
(1977); Raich and Schlesinger (1992). Soil CO2 efflux in the
1-year litter removal plots decreased much faster than the
mean annual decrease during the 7-year continuous litter
exclusion, indicating that the effect of litter input on soil
respiration was nonlinear with a greater reduction at the
earlier stage of litter exclusion.
Root exclusion had a greater effect on reducing soil CO2
efflux in the secondary forest than in the plantation,
suggesting more roots living in the secondary forest than in
the plantation. This conclusion was independently
confirmed by the observation of Lugo (1992) and our
2113
Fig. 3. Total microbial biomass (mg C kgK1 soil) at a depth of 25 cm of
soils in the plantation and the secondary forest in the wet (August) and dry
(March) seasons. Common letters on the bars indicate no significant
difference between the treatments at 95% confidence level.
root biomass (meanGSE) data that showed the root
density was 56.3G4.4 g mK2 (root diameter O5 mm)
and 24.6G3.2 g mK2 (!5 mm) in the secondary forest and
44.2G5.4 g mK2 (O5 mm) and 15.6G2.5 g mK2 (!5 mm)
in the plantation (Li et al., 2004). Our observation that roots
contribute total soil CO2 efflux in the secondary forest (70%)
and in the plantation (56%) does not necessarily mean roots
could contribute 70 and 56% to soil CO2 efflux if root were
present because the reduction would partially be attributed to
the loss of soil microbes and soil fauna. These values, which
might be over-estimated, were lower than the
estimated contribution of root and root litter to total soil
CO2 efflux (70–80%) cross a wide range of forests by
Bowden et al. (1993); Nadelhoffer and Raich (1992). This
difference could be explained by the more prolonged
trenching period in our study (7-y) than theirs’ (9 months).
Varied reduction of microbial biomass during the litter
and root exclusion and different responses of the microbial
biomass to seasonal changes between the plantation and the
secondary forest suggest the microbe-driven decomposition
is ecosystem/vegetation type dependent. Moreover,
microbial activity in the plantation may be more sensitive
to the litter input than the secondary forest because soil
2114 Y. Li et al. / Soil Biology & Biochemistry 36 (2004) 2111–2114
respiration decreased more significantly in the litter exclu-
sion plots than in the control plots.
Acknowledgements
This study was supported by the Luquillo Long-Term
Ecological Research (LTER) and National Science Foun-
dation. We thank Professor Xiaoming Zou for reading
through the previous version of the manuscript and
providing valuable comments. We also thank Joseph Paulin
for checking the language of the revised manuscript,
especially the grammar of the text.
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