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Biomass and productivity of fine and coarse roots in five tropical mountain
forests stands along an altitudinal transect in southern Ecuador
Gerald Mosera; Christoph Leuschnera; Marina Rdersteina; Sophie Graefea; Nathalie Soetheb; Dietrich
Hertela
a
Plant Ecology, Albrecht-von-Haller Institute for Plant Sciences, University of Gttingen, Untere
Karsple 2, 37073 Gttingen, Germany b Plant Nutrition and Fertilization, Humboldt University of
Berlin, Invalidenstrae 42, Berlin, Germany
Accepted uncorrected manuscript posted online: 27 August 2010
Online publication date: 27 August 2010
To cite this Article Moser, Gerald , Leuschner, Christoph , Rderstein, Marina , Graefe, Sophie , Soethe, Nathalie and
Hertel, Dietrich(2010) 'Biomass and productivity of fine and coarse roots in five tropical mountain forests stands along
an altitudinal transect in southern Ecuador', Plant Ecology & Diversity, 3: 2, 151 164, doi:
10.1080/17550874.2010.517788, First posted on: 27 August 2010 (iFirst)
To link to this Article: DOI: 10.1080/17550874.2010.517788
URL: http://dx.doi.org/10.1080/17550874.2010.517788
Biomass and productivity of fine and coarse roots in five tropical mountain forests stands
along an altitudinal transect in southern Ecuador
TPED
Gerald Mosera, Christoph Leuschnera*, Marina Rdersteina, Sophie Graefea, Nathalie Soetheb and Dietrich Hertela
Root production in tropical forests
aPlant Ecology, Albrecht-von-Haller Institute for Plant Sciences, University of Gttingen, Untere Karsple 2, 37073 Gttingen,
Germany; bPlant Nutrition and Fertilization, Humboldt University of Berlin, Invalidenstrae 42, 10115 Berlin, Germany
Introduction
Few studies in the past have addressed the contribution
of root systems to the productivity of tropical forest ecosystems, and when they have done, methodological comparisons have dominated the discussion. A variety of
different approaches to determine fine root production
have been used, the most common being the ingrowth
core and the sequential soil coring methods, recently
supplemented by the mini-rhizotron technique. Reflections on the growth rate of coarse and large roots in tropical forests have been entirely hypothetical (Clark et al.
2001a). As a result the below-ground production has
played a minor role in most publications on net primary
production in tropical forests, or only rough estimates
have been given.
The change in productivity of tropical moist forests
along altitudinal transects has been studied in a number of
regions, including Malaysia (Kitayama and Aiba 2002),
Puerto Rico (Weaver and Murphy 1990; Wang et al. 2003)
and Hawaii (Raich et al. 1997). However, in all of these
studies only above-ground net primary production (ANPP)
has been assessed. Since empirical data based on measurements in tropical montane forests are few, below-ground
productivity has so far been included only as a constant
portion of ANPP (Wang et al. 2003).
A review by Clark et al. (2001b) gave a range for
below-ground net primary production (BNPP) in tropical forests of 1.7 to 21.7 Mg C ha1 year1 (20 to 120% of ANPP),
*Corresponding author. Email: cleusch@gwdg.de
ISSN 1755-0874 print/ISSN 1755-1668 online
2010 Botanical Society of Scotland and Taylor & Francis
DOI: 10.1080/17550874.2010.517788
http://www.informaworld.com
which demonstrates the importance of BNPP in productivity estimates. Such a wide range is a reflection of the fact
that our knowledge on root production in tropical forest
ecosystems is very limited, and an adequate model is not
yet available to predict the ratio of above- to below-ground
production along environmental gradients such as precipitation, temperature and nutrient availability. Until
recently, the lack of precise data on root production in
tropical montane forests has generated considerable uncertainties about carbon (C) balance and turnover in these
forests.
We have conducted a comprehensive study on tree
root mass and root dynamics in five tropical mountain
forest stands in southern Ecuador, firstly to quantify the
biomass of fine and coarse roots along an altitudinal
transect from 1000 to 3000 m elevation above sea level
(a.s.l), secondly to compare estimates of fine and coarse
root production by using three independent methods, and
finally to investigate the influence of elevation-associated
changes in environmental factors on root biomass and root
production.
Material and methods
Study sites
The study was conducted on the eastern slopes of the south
Ecuadorian Andes, about 500 km south of the Equator.
Five forest stands, at 1050, 1540, 1890, 2380 and 3060 m
152
G. Moser et al.
Table 1. Percentage contribution of plant families to the total number of stems in the five study plots along an altitudinal transect from
1050 m (Stand 1) to 3060 m (Stand 5) elevation in southern Ecuador.
Plant families
Melastomataceae
Myrtaceae
Lauraceae
Araliaceae
Rubiaceae
Chloranthaceae
Styracaceae
Ericaceae
Cunoniaceae
Symplocaceae
Aquifoliaceae
Clusiaceae
Cyrillaceae
Gentianaceae
Lecythidaceae
Podocarpaceae
Myrsinaceae
Euphorbiaceae
Monimiaceae
Sapindaceae
Clethraceae
Alzatheaceae
Dubiaceae
Arecaceae
Burseraceae
Theaceae
Celastraceae
Annonaceae
Cecropiaceae
Moraceae
Mimosaceae
Sapotaceae
Myristicaceae
Not determined
Stand 1
Stand 2
Stand 3
2.5
6.3
2.5
1.3
1.3
40
1.3
11.3
31.3
3.8
2.5
1.3
1.3
7.5
2.5
5
5
8.8
1.3
54.7
1.3
5
3.8
1.3
1.3
3.8
23.3
12.5
2.5
5
2.5
2.5
1.3
1.3
7.5
3.8
5
5
1.3
3.8
Stand 4
16.3
1.3
3.8
7.5
3.8
1.3
1.3
8.8
6.3
16.3
1.3
1.3
10
1.3
Stand 5
5
2.5
1.3
15
2.5
2.5
7.5
23.8
5
7.5
10
Important genera
Axinea, Graffenrieda, Miconia
Ocotea, Aniba
Schefflera
Cinchona, Palicourea
Hedyosmum
Styrax
Weinmannia
Symplocus
Ilex
Clusia
Purdiaea nutans
Macrocarpaea
Eschweilera
Podocarpus
Myrsine
Alchornea
Siparuna
Matayba
Clethra
Alzathea
Euterpe
Protium
Maytenus
Guatteria
Cecropia
Ficus
Inga
Proteria
14.7
19.4
17.4
A total of 80 trees were surveyed per plot; species determination were carried out by J. Homeier, University of Gttingen.
153
Table 2. Location data and physiographic characteristics (mean, range) for the five stands in southern Ecuador.
Elevation (m)
Coordinates
Slope ()
Temperature (C):
Air
Organic layer
Mineral soil (10 cm)
Rainfall (mm year1)
Soil type
Soil water content (vol%):
Organic layer
Mineral soil
Organic layer depth (mm)
pH (CaCl2) (Ah)
C:N (L/Of1)
Base saturation (Ah) (%)
Stand 1
Stand 2
Stand 3
Stand 4
Stand 5
1050
S 04 06 54
W 78 58 02
26
1540
S 04 06 42
W 78 58 20
10
1890
S 03 58 345
W 79 04 648
31
2380
S 03 59 19
W 79 04 55
28
3060
S 04 06 71
W 79 10 581
27
9.9 (4.416.0)
29.7 (15.338.5)
48
3.9
22
12.5
12.9 (3.423.9)
30.3 (20.443.5)
243
3.9
29
7.5
11.6 (3.622.3)
35.4 (27.444.7)
305
3.5
28
6.8
34.0 (23.839.8)
44.7 (35.748.7)
214
3.3
46
6.8
45.3 (30.561.7)
49.1 (39.559.5)
435
2.9
63
22.7
Air temperature (at 1.5 m height inside the stands) and soil temperature (annual mean and range in parentheses) were measured between May 2003 and
April 2004. Rainfall was recorded in gaps at 1050, 1950, 2680 and 3170 m and extrapolated to the study stands (data from P. Emck, University of Erlangen: Stands 35, and our own measurements). Volumetric soil water content was measured continuously using TDR probes in the organic layer and the
mineral soil at a depth of 10 cm (annual means and extremes). pH (CaCl2) of the mineral topsoil (030 cm), C:N ratio of organic layer (L/Of1), base saturation (Ah) and soil type were determined by Iost (2007).
Table 3. Above-ground forest structure of the five stands studied. Values are means, SE where applicable.
Stand 1
Stand 2
Stand 3
Stand 4
Stand 5
31.8
6.0 0.4a
17.3 1.3a
33.6
21.7
5.4 0.4a
11.5 0.6b
27.5
18.9
5.7 0.5a
12.2 0.8b
36.9
12.0
2.8 0.2b
9.8 0.6c
27.2
9.0
2.2 0.2c
7.2 0.4d
42.2
Canopy height, DBH and basal area are from Moser et al. (2008); LAI from Moser et al. (2007). Different letters indicate statistically significant differences
between the stands (P < 0.05).
154
G. Moser et al.
Ingrowth cores
In April 2003 an ingrowth core study was established. In
each of the five stands, 20 soil cores (55 mm diameter and
25 cm depth) were taken from the randomly distributed
subplots used for the sequential coring approach. All visible fine roots in the soil were carefully removed by hand
and the soil material was reinserted into the holes and
compressed to about the original soil density. Holes were
filled by adding soil from additional cores, thus replacing
the volume of the roots extracted. Organic layer and mineral topsoil samples were treated separately. The cleaned
material was filled back into the holes to re-establish reasonably natural soil conditions. The ingrowth cores were
marked with PVC tubes and recovered after 15 months
using the same soil corer as used originally. The soil was
separated into organic and mineral fractions and placed in
polyethylene bags. To estimate the impact on fine root
growth caused by root injury during core removal, 10
additional ingrowth cores were taken per plot. Two of
these were re-sampled in each plot after 6, 12, 18, 24 or 30
weeks and the fine root biomass ingrowth determined.
In the ECSF laboratory the soil samples were washed
and all the fine rootlets (> 10 mm length) were extracted
by hand, washed and dried (70 C, 48 h). The study with
sequential re-sampling of ingrowth cores indicated that
fine root ingrowth into the cores began about three months
after the start of the experiment. It was thus assumed that
the fine root biomass detected in the cores after 15 months
represented a fine root growth period of 12 months. As the
depth of the ingrowth cores was 25 cm, production estimates obtained using this method applied only to the
organic layer in the high-elevation stands, which had thick
humus layers. Nevertheless, with the fine root biomass
data from the organic layer and the mineral topsoil, the
ingrowth core production estimates could be extrapolated
to the soil profile covered by the sequential coring
approach (organic layer plus mineral soil to 30 cm).
Mini-rhizotrons
As a further independent method for studying fine root
dynamics, 10 transparent mini-rhizotron tubes with an
external diameter of 70 mm were installed in each stand in
June 2005. These were placed perpendicular to the soil
surface and wherever possible inserted at a depth of about
40 cm. Only the first 10 cm below the soil surface was
considered for the assessment of fine root dynamics, as in
a number of cases the very high stone content of the mineral soil at the lowermost stand did not allow the tubes to
be installed more deeply. To monitor root growth, a root
scanning system (CI600 Root Growth Monitoring System,
CID Inc., USA) was employed at monthly intervals until
155
January 2007. In order to avoid artefacts caused by disturbance during installation of the tubes, only the data from
November 2005 onwards were considered for analysis.
The images were analysed using the programme WinRHIZO
Tron (Rgent, Quebec, Canada). Only fine roots were used
for the analysis. Relative root length production and relative root loss were calculated by relating root length production or root loss between two observation dates to the
root length visible at the previous measuring date. Extrapolation of these data to a full year allowed estimation of
average annual root production and loss rates, root turnover rate as a percentage of the standing stock, and root longevity (days) of the visible fine root population in the
rhizotron tubes (Nadelhoffer 2000). From turnover rates in
the uppermost 10 cm and biomass data from the organic
layer and mineral topsoil (030 cm), annual fine root production was calculated for the five stands.
Fine root litter decomposition
Decomposition of fine root litter was studied using two
different litter bag studies, largely following the protocol
proposed by Fahey et al. (1988). In the first of the studies
we investigated the decomposition rate of dead fine root
material at the site of origin. During March 2003 live fine
root material was sampled from the organic layer of each
stand, cleaned of soil residues and 5 g fresh weight of this
material inserted into litter bags (nylon, 10 cm 10 cm,
mesh 1 mm, n = 12). Additional fine root material was
immediately dried for determination of water content. The
litter bags were exposed within the organic layer at random locations within each plot and covered by a thin layer
of leaf litter. After 12 months exposure in the field, the
litter bags were recovered, and the remaining undecomposed roots carefully cleaned of soil residues, dried and
weighed. In a second litter bag study, starting January 2004,
we sampled live fine roots from the mid-elevation Stand 3
(1890 m a.s.l.) and exposed this material in the five stands
for six months to estimate the decomposition rate in a
standardised manner, independent of differences in root
properties among the five stands. In each study the decomposition rate was calculated from the difference between
the initial and the remaining root mass in the litter bag.
To obtain information on root chemistry, which determines the ability of fine roots to decompose, the C:N ratio
of fresh fine root biomass to necromass (n = 40) was
measured using a CNAnalyser (Vario EL III, Fa. Elementar, Hanau, Germany) and the polyphenol concentration
in fine roots (n = 10) from the organic layer was determined by photometry (DU 640 Spektralphotometer, Fa.
Beckmann, Germany, data provided by M. Unger). Cleaned
fresh fine root material for the analysis was frozen at
20 C. After the samples had been transported to Gttingen under continuous cooling they were ground to powder
under liquid nitrogen in a mortar. For the assessment of
soluble and insoluble tannin concentration, a repeat
procedure was conducted by suspending the sample in
methanol (twice) and subsequently in acidbutanol and
156
G. Moser et al.
calculations were carried out using Xact software (SciLab,
Hamburg, Germany, version 8.0).
Results
Root biomass distribution
Total below-ground biomass (fine, coarse and large roots)
was largest at the highest elevation stand, both coarse and
large root biomass being significantly higher than those in
the lower stands. Between 1050 and 2380 m a.s.l., total
estimated root biomass ranged from 26.1 to 39.4 Mg ha1
(Table 4).
The increase in total root biomass with elevation was
mainly due to a conspicuous increase in root mass within
the organic layer, which itself increased about nine-fold in
thickness. Most of the root biomass in Stands 25 was
concentrated in the organic layer, reaching 83% of the
total profile in Stand 4 and 77% in Stand 5. Below a depth
of 30 cm in the mineral soil only a very few coarse and
large roots were found. Total root biomass correlated with
all seven environmental factors tested, but the closest
relationship existed between root mass and soil C:N ratio
and root mass and soil proton concentration (Table 5).
The contribution of fine root to total root biomass
increased with elevation. Similarly to total root mass, fine
root biomass also correlated significantly with all the
environmental parameters tested, apart from precipitation.
Fine root biomass increased linearly with elevation, while
necromass increased exponentially (Figure 1). In the two
lowermost stands, fine root biomass and necromass were
similar; in contrast, at the highest altitude necromass
exceeded biomass by a factor of 2.7. Coarse and large root
biomass showed a high spatial variability among the
stands, particularly in the lowermost stands, which may
Statistical analysis
Differences in root biomass, root production and decomposition rates among the five stands were analysed by the
KruskalWallis test and the MannWhitney two-sample
test (U-test) using the package SAS (SAS Institute, Cary,
NC, USA, version 8.2).
Linear and simple non-linear regression analysis was
used to identify the significance of elevation, mean air
temperature, annual precipitation, soil moisture and proton
concentration in the mineral soil, or the C:N ratio of the
organic layer, on root biomass and root production
estimates. Additional regression analysis was performed
on the relationship between the polyphenol content or
C:N ratio of fine roots and litter decomposition rate. The
Table 4. Means SE of standing biomass of coarse and large roots (diameter > 2 mm) and living fine roots (< 2 mm) from soil coring
in the organic layer (variable in depth), mineral soil (30 cm) and the total profile to 30 cm depth in Mg ha1 in the five forest stands.
Coarse and large roots n = 12; fine roots n = 15.
Stand 1
Coarse + large roots
Stand 3
4.0b
3.0a
8.2a
0.3b
0.2a
0.4b
Stand 4
2.1b
1.7b
2.6b
0.2c
0.2b
0.3b
22.2
8.1
30.7
3.6
2.2
5.6
36.3
2.2
26.8
29.4
0.5
2.2
2.7
32.1
organic
mineral
profile total
organic
mineral
profile total
profile total
Fine roots
Stand 2
0.6a
7.5a
8.4ab
0.1a
0.2a
0.3a
Stand 5
4.9b
0.6b
5.7a
0.3b
0.3ab
0.6b
12.3
4.7
19.6
2.4
3.3
6.2
26.1
37.8 4.3b
8.5 2.3a
51.9 4.7c
6.5 0.3d
4.8 0.3c
10.8 0.6c
62.7
24.0
3.0
32.7
3.6
2.8
6.3
39.4
Table 5. Results of correlation analyses between three root biomass fractions and stand climatic and edaphic variables.
Elevation
Dependent
r2
r2adj
Temperature
P
r2
r2adj
Rainfall
r2
r2adj
Soil moisture
P
r2
r2adj
Soil acidity
r2
r2adj
C:N ratio
r2
r2adj
Coarse + large roots 0.35 0.13 0.149 0.39 0.18 0.131 0.35 0.14 0.145 0.34 0.12 0.153 0.61 0.48 0.059 0.59 0.46 0.064
Fine roots
0.90 0.87 0.006 0.89 0.86 0.007 0.37 0.16 0.140 0.71 0.61 0.036 0.85 0.80 0.012 0.83 0.77 0.015
Total root mass
0.69 0.58 0.040 0.73 0.64 0.032 0.68 0.57 0.043 0.91 0.82 0.043# 0.84 0.78 0.014 0.90 0.87 0.006
Statistically significant results are printed in bold (P < 0.05); #, nonlinear relationships.
80
Necromass
3000
70
Mass loss per year (%)
Elevation (m a.s.I.)
157
2500
2000
1500
Standard material
60
50
40
30
Local material
20
10
1000
0
1000
3000
3000
2000
1000
0
1000
2000
Fine root biomass (g m2) Fine root necromass (g m2)
3000
1500
2000
2500
Elevation (m a.s.l)
partly bias the overall stand-level estimates of root biomass in the study. The largest quantities of coarse and
large root biomass were present in Stand 5 (52 Mg ha1),
and the lowest in Stand 3 (20 Mg ha1, Table 4). The biomass of large diameter roots showed no significant relationship with any of the environmental variables examined
(Table 5).
Fine root decomposition
The decomposition rate of fresh fine root material when
exposed at the site of origin was very similar at the three
lowermost elevations, but decreased in the two uppermost
stands to about a third of the rate of that at 1050 m (Figure 2).
The potential decomposition rate, determined by exposing
standard material from Stand 3, was higher in all stands
(except Stand 3) than the decomposition rate of the local
root material. The decomposition of standard root material
showed a trend towards a decrease (P = 0.07) with
elevation.
The concentration of soluble phenols in fine roots
increased significantly with elevation (Table 6), as did the
concentration of insoluble phenols. The concentration of
Root production
The annual production of coarse and large roots (> 2 mm)
was estimated at about 0.2 Mg ha1 year1 in Stands 1, 3
and 4, and was significantly higher in Stands 2 and 5
(Table 7). None of the five environmental variables investigated, nor elevation, showed correlation with coarse and
large root production.
Fine root production, estimated by sequential coring
combined with the maximumminimum calculation procedure, increased more than ten-fold along the transect
Table 6. Mean + SE polyphenol concentration of fine root biomass and C:N ratio of fine root bio- and necromass of the organic layer
in the soil of five forest stands.
g1
Polyphenols (mg
fresh biomass)
PhenolSoluble
PhenolInsoluble
TanninSoluble
TanninInsoluble
C:N ratio
Fine root biomass
Fine root necromass
Stand 1
Stand 2
Stand 3
Stand 4
Stand 5
1.7 0.4a
4.2 0.8a
4.8 1.9a
50.4 8.2ab
2.3 0.3a
12.9 2.3b
6.9 1.1a
57.3 5.6b
2.3 0.5a
7.4 1.6ab
4.9 1.3a
51.0 7.2ab
3.9 1.2b
7.6 1.8ab
6.0 1.0a
34.4 4.2a
4.9 0.7b
19.8 2.6c
14.5 1.7b
45.8 7.2ab
28.2 1.0a
23.6 1.9a
34.2 0.6b
26.9 0.5a
43.4 0.9c
33.1 0.8b
55.2 1.2d
44.2 1.0c
91.1 3.3e
60.1 1.5d
Polyphenols: n = 10; C:N, n = 40; different letters indicate statistically significant differences between the forest stands (P < 0.05).
158
G. Moser et al.
Table 7. Mean + SE of stand level estimates of coarse and large (> 2 mm) root growth in the organic layer and the mineral soil
(050 cm) of five stands, extrapolated from increment rates of 20 roots per stand and coarse and large root biomass data (in Mg ha1).
Organic
Mineral
profile total
Stand 1
Stand 2
Stand 3
Stand 4
Stand 5
0.01 0.003a
0.16 0.04
0.17 0.04A
0.58 0.08b
0.21 0.06
0.79 0.14B
0.16 0.02c
0.06 0.02
0.22 0.03A
0.20 0.04c
0.03 0.01
0.23 0.04A
0.72 0.05b
0.16 0.03
0.89 0.08B
Different letters indicate statistically significant differences between the forest stands (P < 0.05).
Table 8. Means + SE of biomass production of fine roots (Mg ha1 year1) in the organic layer and the mineral soil (030 cm) as
estimated with three different methods (and two different calculation procedures in the sequential coring approach).
Soil
Sequential coring
(Comp. flow)
Sequential coring
(Maxmin)
Ingrowth cores
Mini-rhizotrons
Soil horizon
organic
mineral
profile total
organic
mineral
profile total
organic
mineral
profile total
organic
mineral
profile total
Stand 1
0.99
1.17
2.15
1.44
0.50
1.94
0.76
2.86
3.62
0.41
1.87
2.28
0.10a
0.07a
0.13a
0.14a
0.06a
0.10a
0.13a
0.51ab
0.65a
0.04a
0.17a
0.20a
Stand 2
2.72
1.00
3.72
4.79
1.30
6.09
6.08
2.98
9.06
1.95
1.22
3.16
0.21b
0.14a
0.30b
0.50b
0.15b
0.33b
0.67b
0.32b
1.00b
0.31bc
0.20a
0.51ab
Stand 3
14.49
7.70
22.19
4.52
2.24
6.76
3.38
3.64
7.02
1.16
1.58
2.97
Different letters indicate statistically significant differences between the forest stands (P < 0.05).
0.56c
0.21b
0.46c
0.51b
0.24c
0.38b
0.58c
0.62b
1.21bc
0.13b
0.22a
0.31ab
Stand 4
21.14
14.64
35.78
7.89
4.61
12.05
3.55
2.55
6.10
2.12
1.63
3.72
0.95d
0.30c
0.97d
0.99c
0.55d
0.77c
0.71bc
0.51ab
1.23c
0.28c
0.22a
0.50b
Stand 5
32.98 0.93e
42.62 1.68d
75.60 2.14e
10.87 1.42d
11.06 1.27e
21.93 2.15d
2.26 0.29c
1.52 0.20a
3.78 0.49a
5.71 0.76d
4.26 0.57b
9.40 1.32c
Ing Cor
Minirhiz
BNPPMaxMin
Ing Cor
Minirhiz
159
30
Max-Min
25
BNPP (Mg ha1 yr1)
Coarse and large root production was determined by diameter increment measurement at 20 root segments per plot. Fine root production was estimated by sequential coring combined with a maximumminimum
calculation (n = 20 coring locations), ingrowth cores (n = 20) and the mini-rhizotron technique (n = 10); elevation, m a.s.l., mean air temperature in C, annual precipitation in mm year1, mean soil moisture of mineral
soil in vol%, soil acidity refers to proton concentration in solution from the mineral soil (in mmol l1), C:N ratio refers to the organic layer. r2 and r2adj refer to linear or simple nonlinear regression models. Significant
results are printed in bold (P < 0.05), non-linear relationships are marked by #.
0.132
0.001
0.280
0.004#
0.007
0.344
0.010#
0.18
0.97
0.17
0.98
0.86
0.26
0.95
0.38
0.98
0.13
0.99
0.90
0.06
0.98
0.19
0.94
0.01
0.94
0.67
0.14
0.89
0.14
0.91
0.14
0.98
0.62
0.20
0.95
0.15
0.96
0.15
0.99
0.72
0.10
0.97
0.331
0.059
0.291
0.134
0.059
0.322
0.147
0.24
0.48
0.18
0.17
0.48
0.23
0.13
0.07
0.61
0.11
0.38
0.61
0.08
0.35
0.168
0.006#
0.320
0.003#
0.016
0.377
0.010#
0.07
0.97
0.22
0.98
0.77
0.29
0.95
0.30
0.98
0.08
0.99
0.82
0.04
0.97
0.168
0.006#
0.358
0.003#
0.015
0.415
0.010#
0.30
0.98
0.05
0.99
0.83
0.02
0.97
Dependent variable
0.07
0.97
0.27
0.98
0.78
0.31
0.95
P
adj
P
adj
P
adj
0.236
0.018#
0.238
0.005#
0.035
0.305
0.011#
0.39
0.95
0.25
0.95
0.75
0.15
0.92
0.129
0.002
0.199
0.002
0.028
0.236
0.004
r2adj
adj
r2
Soil moisture
r2
r2
Rainfall
r2
r2
Temperature
r2
Elevation
r2
r2
Table 9. Correlation analyses between three root production fractions and six climatic and edaphic variables.
adj
r2
Soil acidity
r2
r2
C:N ratio
20
15
10
Ing Cor
Minirhiz
5
0
1000
1500
2000
2500
Elevation (m a.s.l.)
3000
160
G. Moser et al.
hypothesis advanced by Northup et al. (1995), the complexing of nitrate ions by polyphenols can reduce the loss
of these ions with drainage water from N-deficient soils,
thereby increasing the concentration of dissolved organic
N, but reducing that of NH4+ and NO3. Thus, elevated
phenolic concentrations in soil water may conserve N, and
can also ameliorate Al3+ toxicity through complexing (Hue
et al. 1986). In general, nitrification in many soils is
reduced at pH values < 5.5, but on the other hand nitrifying
fungi may produce NO3 in acidic soils. Krashevska et al.
(2008) showed for our transect that the microbial community
changed with elevation from being bacteria-dominated at
1000 m to fungal-dominated at 3000 m. Despite the low
pH of the mineral topsoil in all five stands, the nitrate and
ammonium concentrations in Stands 13 were quite high
and decreased significantly towards the uppermost stands,
4 and 5 (S. Iost, pers. comm.). It therefore seems unlikely
that low pH was a key factor in limiting nitrification in the
upper section of our transect, and more likely that other
factors such as decreases in litter quality and temperature,
and increasing soil moisture and soil phenol concentrations, were responsible for the decrease in N. It was
assumed by Hafkenscheid (2000) that the large amounts of
phenols in the topsoil water of stunted Jamaican mountain
forests were probably exudates from fine roots rather than
leachates from leaf litter. In our own transect, we found a
more or less constant concentration of soluble phenols in
the leaves (M. Unger, pers. comm.), but an increasing concentration of fine root mass with elevation which may support Hafkenscheids hypothesis.
We hypothesise that adverse soil conditions in the
upper montane forests, in particular low N and probable
waterlogging, and possibly also high concentrations of
Al3+ and other toxic cations, may have restricted the nutrient supply, thereby stimulating the trees to enlarge their
fine root system and release significant amounts of
polyphenols through root exudation. Increased production
of secondary metabolites in the root system, together with
high fine root production, which compensates for root
losses in an adverse environment, should represent a notable sink for C and nutrients, which in turn must reduce
above-ground productivity and tree height. In the case of
coarse and large roots, strong winds on exposed ridges and
at high elevations may be an additional factor in southern
Ecuador that shifts C allocation to the root system.
Fine root production measurement in tropical
forests an assessment of methods
Application of the three methods for estimating fine root
production has, in the past, produced highly divergent
results (Nadelhoffer and Raich 1992; Publicover and Vogt
1993; Fahey et al. 1999; Hertel and Leuschner 2002;
Hendricks et al. 2006). This has been confirmed in our own
study, and supports the need for a critical evaluation when
these are applied to tropical montane forests.
The sequential coring approach. Estimates of fine root
production by soil coring are dependent on accurate periodic
Seasonal CV organic
mineral
Spatial CV
organic
mineral
Stand Stand
1
2
Stand
3
Stand Stand
4
5
0.81
0.33
0.61
0.32
1.10
1.09
0.35
0.56
1.06
1.17
0.32
0.48
0.30
0.55
0.15
0.57
1.14
1.38
0.55
0.70
161
have masked this effect, leading to a significant underestimate of fine root production in Stands 1 and 2. At higher
elevations, with cool soil temperatures and frequent waterlogging, the delay period may have been longer than three
months, causing an underestimation of root production at
2340 and 3060 m.
Mini-rhizotron approach. This technique enables the
direct observation of fine root growth and death in the
rhizosphere. When combined with root biomass determination, this approach may yield more reliable estimates of
fine root production of forests than any other available
technique (Hendricks et al. 2006), particularly in climates
with low seasonality, like Ecuador. The installation of the
rhizotron tubes may represent a similar disturbance to the
rhizosphere as that with the ingrowth core approach. It
requires a recovery period of at least six months before
root initiation and death may have regained a quasi-steady
state again. In cool environments with slower root growth
(e.g. in the present study at 2450 m and 3060 m elevation),
recovery may take even longer (Graefe et al. 2008). Minirhizotron studies along temperature gradients may thus
encounter different recovery periods at the sites being
compared. To counter this, we focused on root length loss
rather than root growth, and related this to mean standing
fine root biomass in calculating root turnover. Even if new
root growth partly represented a compensation for losses
due to the initial disturbance event, root death would
reflect mortality caused by physiological and local environmental factors, thus giving a better estimate of turnover
than calculations based on root growth.
Estimates of fine and coarse root production
With the mini-rhizotron approach, we estimated root
production in the four lowermost stands (14) to be 2.3 to
3.7 Mg ha1 year1, indicating a significant increase along
the elevation transect when Stand 5 was also taken into
account. However, the very high value obtained for Stand
5 at 3060 m (9.4 Mg ha1 year1) may represent an overestimate, for two possible reasons. Firstly, it is unclear if
the root growth and death events observed in the minirhizotron tubes in the uppermost 10 cm of the soil profile
were representative of the entire profile, in particular in
the very thick, acid and moist organic soils of the uppermost stands. It is also questionable if the growth pattern of
the few visible fine roots in the tubes at 3060 m were representative of all roots growing in the uppermost 10 cm of
this soil profile. This type of error should be less severe in
the warmer soils with thinner organic layers at lower elevations, where the number of roots observed was much
higher. Secondly, it is likely that fine root growth would
be slower at higher elevations with lower temperatures,
and this would suggest that longer equilibration periods
should be allowed for the mini-rhizotron tubes after
installation in high-elevation forests (Graefe et al. 2008).
Since we had no data on the duration of the installation
shock at different elevations, we used an identical putative
equilibration period (six months) throughout the transect.
162
G. Moser et al.
Notes on contributors
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