ISSN 10642293, Eurasian Soil Science, 2010, Vol. 43, No. 6, pp. 693–700. © Pleiades Publishing, Ltd., 2010. Original Russian Text © O.V. Masyagina, S.G. Prokushkin, T. Koike, 2010, published in Pochvovedenie, 2010, No. 6, pp. 740–747.
DEGRADATION, REHABILITATION, AND CONSERVATION OF SOILS
The Influence of Thinning on the Ecological Conditions and Soil Respiration in a Larch Forest on Hokkaido Island O. V. Masyaginaa, b, S. G. Prokushkina, and T. Koikeb a
Sukachev Institute of Forestry, Siberian Branch, Russian Academy of Sciences, Akademgorodok 50, Krasnoyarsk, 660036 Russia b Hokkaido University, Sapporo, 0608589 Japan Email:
[email protected] Received March 6, 2008
Abstract—The effects of cutting on the ecological conditions and soil respiration in larch forests of Japan were studied. The cutting was found to significantly change the soil surface, resulting in the high spatial and temporal variation of the hydrothermal conditions and soil respiration. The influence of a stand’s thinning on the environment and soil respiration is considered using the example of the thinning of a ripening larch stand in the Tomakomai National Forest (Hokkaido Island, Japan). The changes in the hydrothermal condi tions (the temperature and moisture of the mineral soil layers and litter) and some other factors (the root and litter density and the C/N ratio) after the thinning of the stands and their influence on the soil respiration were studied. The soil respiration in the thinned forest site did not differ from that on the control plot, whereas the soil temperature was much higher in the former. The moisture of the soil mineral layers and the litter was the same. Despite the latter fact, no significant relationships between the soil respiration and its temperature and moisture were found. In the area covered with the thinned forest, the water content of the litter turned out to be the main microclimatic factor affecting the soil respiration. There, the fine roots and litter density were 18 and 15% less, respectively. The thinning of the stand induced high variation of the soil respiration and tem perature, as well as of the fine roots and the litter density. On the whole, the soil respiration in the larch forest studied in Japan was determined by the litter stock and the C to N ratio. DOI: 10.1134/S1064229310060104
INTRODUCTION The soil respiration (Rs) is a term characterizing the processes related to the CO2 emission from the soil surface as a result of the root respiration and the oxi dation of detritus and organic matter by the soil micro flora. Their intensity depends on the temperature (Ts) and water content of the soil mineral layer (MSWC) and the litter (LWC), the edaphic and phytocenotic conditions, and the specificity of the distribution of assimilates in the biomass of the phytocenosis [4]. The temporal and spatial variation of the soil respiration is a result of the variability of the ecological conditions [3]. Therefore, studies of these factors, along with the determination of the carbon content and the root bio mass, promote the identification of the causes respon sible for the variation of the soil respiration [2]. At present, the data obtained in similar investigations of humanmodified ecosystems are insufficient for esti mating the role of this process on the global scale. Cuttings, including improving ones, differently affect the carbon cycle in forest ecosystems. First, tree biomass—the main component of forest ecosystems [1, 13]—is removed as a result of cutting; second, the removal of the photosynthetic apparatus decreases the degree of carbon assimilation by the whole phyto
cenosis immediately after cutting. The latter circum stance affects the productivity of forest stands. Lastly, the improvement cuttings change the physicochemi cal properties of the soil and, hence, of the fluxes of carbon, nitrogen, and ash elements. Particularly, they lead to an increase in the air and soil temperature [5, 16], whereas the reduction in the total transpira tion of the cenosis raises the soil moisture [9]. Since the soil respiration is related to the soil temperature and moisture [4, 11], their increase enhances the CO2 emission from the soil. The effect of the improvement cuttings on the availability of mineral elements and the microbial activity was also revealed [10]. The influ ence of partial harvesting on the CO2 emission from the soil of forest ecosystems has been poorly studied in Russia. Only in foreign literature is there information on the effect of improvement cuttings on the soil res piration. The thinning of stands was shown to intensify the soil respiration due to the activation of metabolic processes related to the detritus and organic matter pools [6, 14, 17, 21]. The decrease in the respiration activity might be due to the inhibited root respiration as the roots die off after the cuttings [23]. The com bined effect of the cuttings and their consequences leads to a decrease in the soil’s carbon pool by 10–50%
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Table 1. Characterization of the climate in the studied region (data of Qu [22]) Parameter
Value
Mean annual precipitation, mm 1250 Mean annual temperature, °C +7.3 Mean August temperature, °C +19.1 Mean January temperature, °C –3.2 Air temperature at a height of 1.3 m from the soil surface from June to October 2003, °C maximal +26.4 minimal +0.8 mean +16.1 ± 0.1 Soil temperature at a depth of 10 cm from June to October 2003, °C maximal +20.9 minimal +9.7 mean +16.3 ± 0.1 Table 2. Characteristics of the stand before (above the lines) and after (under the lines) the thinning in January 2004 (per sonal information of Prof. Hirano) Stand density
The stand aboveground biomass
Basal area
Species ind./ha
% of the sampling
m2/ha
% of the sampling
t/ha
% of the sampling
Larch
716 480
33
24.1 17.7
27
82.8 61.2
26
Broadleaved
505 421
17
5.9 5.4
9
23.8 22.2
7
Mean height of the trees (data of 2000)
Canopy height
Age (data of 2003), years
m 18–20*
8.9*
50*
Not det.
* According to Qu’s data [22].
under the intense use of a forest [14] and changes in the rate of the N2O, CO2, and CH4 exchange between the soil and the atmosphere [7]. The aim of this work is to study the spatial variation of the soil respiration related to the thinning manage ment that change the environment of the stands.
FraserJenk. et Jermy) and Pachysandra terminalis Sieb. et Zucc.
OBJECTS AND METHODS
The soils are of volcanic origin with a weakly acid reaction (pH 5.0–6.0), are oligotrophic, and they have a low humus content (about 0.1%); the litter thickness is 1.0 to 5.0 cm. The roots concentrate in the upper (0–15 cm) soil layer between the litter and the under lying layer of porous pumice. The total root biomass is nearly 13.1 t/ha [22].
The influence of the stand thinning on the intensity of the soil respiration was studied in a 50yearold larch stand in the Tomakomai National Forest (42°44′N, 141°31′E; 115–140 m a.s.l.; Hokkaido Island, Japan). This region is characterized by a humid climate (Table 1). The stand is mainly composed of Japanese larch (Larix kaempferi (Lam.) Carr.) with an admixture of Picea jezoensis Sieb. et Zucc.) and broad leaved species—birch (Betula sp.), oak (Quercus sp.), and magnolia (Magnolia sp.) (Table 2). The under story of the stand mainly consists of ferns (Dryopteris crassirhizoma Nakai. and Dryopteris expansa (C. Presl)
The thinning of the stand was carried out in Janu ary 2004, when the snow cover was of 0.5 m deep and the soil was frozen. Therefore, the soil and the ground cover were almost undisturbed. The wood and cutting debris were removed from the plot. The characteriza tion of the stand before and after the thinning is given in Table 2. In our research, the effect of thinning turned out to be insignificant, since only 33% of the larch trees and 17% of the broadleaved species were removed (Table 2). The data on the effect of the thin ning on the environment and the soil respiration were compared with the same characteristics in an undis EURASIAN SOIL SCIENCE
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THE INFLUENCE OF THINNING ON THE ECOLOGICAL CONDITIONS (a) 20 18 16 14 12 10 8 6 6
EXPERIMENTAL The soil respiration (Rs) was measured during 4 days (July 27–30, 2004) half a year after the thinning of the stand using an LI6400 infrared gas analyzer (LICOR, USA) with an LiCor 600009 soil chamber (LICOR, USA). The measurements were made using fixed soil collars to decrease the errors related to the disturbance of the gaseous soil phase. There were three collars on each plot. The total number of the soil col lars in the control and in the area with the thinned stand was 90 and 109, respectively. The collars with a diameter of 10.2 cm were placed at a depth of 1.5 cm for 10 days before the measurements to equilibrate the soil gaseous phase; they were left there up to the end of the experiment. The vegetation inside the rings was removed. The temperature and soil respiration were measured simultaneously at a depth of 5 cm. The res piration was calculated from the increase in the СО2 concentration in a period of time, the volume of the system (991 cm3), and the area occupied by the soil chamber (71.6 cm2). The measurements were made in the daytime. The average values of the respiration for three soil collars on each plot were used; the variation of the Rs was estimated using the coefficient of varia tion (CV). The significance of the differences between the values obtained was determined using Student’s coefficient. The statistical analysis was performed using the STATISTICA 6.0 StatSoft package. Soil cores in two replicates were taken using an auger within each collar after measuring the soil respi ration to determine the root and litter density, the soil and litter water content, and the C to N ratio. The total of the samples was 180 in the control and 216 on the plot with the thinned stand. The total reserve of roots calculated for the auger volume (the grams of abso lutely dry mass per cubic meter, g a.d.m./m3) and the moisture of the soil and litter were determined using traditional methods. The carbon and nitrogen con tents were measured in dried (at 70°C for 72 h) and sifted soil using an NC analyzer (NC900, Japan). The data were calculated based on the absolutely dry mass. EURASIAN SOIL SCIENCE
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4 4
3 2 1
E
G
I
C
A
(b) 20 18
Soil respiration, mmol/m2/s
turbed stand (control). The size of the plots analyzed was 40 × 40 m. For the assessment of the relationships between the soil respiration and the ecological condi tions, 30 sample plots were arranged in the control area and 36 on the plot exposed to thinning. Their size I was 1 × 1 m. On the same plots, the soil respiration, temperature, and moisture of the mineral soil and litter; the density of litter and fine roots in the 0 to 15cm layer; and the total contents of carbon, nitrogen, and the C/N ratio were determined.
695
16 14 12 10 8 6 6
4 5
4
3
2
1
K
I
G
E
C
A
ber num Plot
Fig. 1. Intensity of the soil respiration at the (a) control and (b) test (thinned stand) plots.
RESULTS AND DISCUSSIONS The soil respiration varied from 5.6–12.1 in the control to 4.5–19.8 μmol/m2/s in the disturbed plot (Fig.1). Its variation was greater on the plot with the thinned forest (CV 33%) than in the control (19%). Despite this fact, no significant differences in the soil respiration between the control and thinned plot were revealed. Such a tendency was noted in the works of many specialists who studied coniferous forest ecosys tems. According to Carter et al. [8], the partial harvest in pine forests (Pinus taeda L.) did not affect the soil respiration and, in both plots, it amounted to about 11 g/m2/24 h (or 3 μmol/m2/s) at 17.5°C. Ohashi et al. [20] found that, for the first year after the thinning of the Cryptomeria japonica D. Don) stand, the soil respira tion in July varied within 442–520 mg/m2/h (or 2.8–
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Table 3. Mean values (±error of the mean) of the soil respiration and the factors affecting it on the control and experimental plots Factor
Control
Soil respiration (Rs), µmol/m2/s Soil temperature (Ts), °C
8.1 ± 0.3 +19.6 ± 0.06
Plot with the thinned stand
Student’s coefficient
8.3 ± 0.5
–0.33
+21.5 ± 0.1
–12.74
P 0.73 0.00000
Water content of the mineral soil layer (MSWC), %
45.6 ± 1.4
47.6 ± 1.2
–1.13
0.26
Litter water content (LWC), %
67.2 ± 0.7
65.3 ± 0.7
1.83
0.07
Density of the fine roots, g/m3
74.0 ± 3.9
60.6 ± 9.0
1.29
0.20
1344.2 ± 44.7
1150.5 ± 49.7
2.86
0.006
C, %
11.5 ± 0.9
12.4 ± 0.7
–0.82
0.42
N, %
0.75 ± 0.03
0.91 ± 0.04
–3.29
0.002
C/N
15.3 ± 1.0
13.5 ± 0.4
3
Litter density, g/m
3.3 μmol/m2/s) and did not differ from that in the control (482 mg/m2/h or 3 μmol/m2/s). In these investigations, the CO2 emission from the soil after the cutting was equivalent to the Rs in the control plots due to some compensation for the reduction in the root respiration by the increase of the microbial respi ration due to the additional input of organic residues after cutting. Probably, this fact is a cause of the absence of significant changes in the soil respiration after the cutting. For the determination of the role of the ecological factors in the changes of the soil respiration rate, mul tiple regression analysis was performed. This analysis revealed that the soil respiration in the control (R 2 = 0.54, F = 5.55, P = 0.002, N = 30) depends on the combined effects of the soil temperature (the par tial correlation coefficient (PCC) = 0.42, P = 0.03), its moisture (PCC = –0.49, P = 0.01), the roots (PCC = 0.27, P = 0.18), and litter density (PCC = –0.24, P = 0.24). On the plot with the thinned stand (R2 = 0.42, F = 7.55, P = 0.0006, N = 36), the soil respiration depended on the soil water content (PCC = –0.48, P = 0.005), the litter density (PCC = 0.43, P = 0.01), and the C to N ratio (PCC = 0.31, P = 0.08). In the control, the combined effect of such factors as the temperature, moisture, and density of roots and litter is responsible for half (54%) of the soil respiration. On the thinned plot, 42% of the variation in the soil respi ration was related to the combined influence of the water content, litter density, and the C/N ratio. Among the microclimatic parameters, the litter mois ture was the most significant; it correlated strongly with the soil respiration. The comparison of the soil respiration rate with that in other regions showed that the values obtained in this experiment were comparable with the results from different ecosystems under similar temperature
1.84
0.07
conditions. According to the estimates of Klopatek [15], the soil respiration in a fir forest (Washington state, USA) was 5 μmol/m2/s. By the data of Yim et al. [26], the mean soil respiration in a larch stand (Hok kaido, Japan) in late August was about 797 mg/m2/h (8 μmol/m2/s). During the period of observations, the temperature and precipitation corresponded to the norms typical for the particular season [22]. The mean temperature of the control soil differed from that in the thinned plot due to the lower density of the stand after the thin ning (Table 3, Fig. 2). The variation of the soil temper ature on the experimental (20.3–22.9°С, CV = 3%) plot and in the control (18.5–20.1°С, CV = 2%) plot was similar (Fig. 3). The thinning did not affect the soil moisture as well. It was close in both plots (Table 3), as well as the range of its variation (25–58% in the control plot, CV = 17%; 26–63% in the test area, CV = 15%) (Fig. 2). The water content of the litter in the plots also did not differ—54–74% (CV = 7%) on the plot with the thinned forest stand and 58–75% (CV = 6%) in the control area (Table 3). Despite the significant differ ences in the soil temperature (Ts) between the plots, the water content of their mineral soil layer (MSWC) and litter (LWC) had the same dynamics on both plots (Fig. 2). Some authors [19, 25] consider that thinning activates the decomposition of organic matter in the soil and, as a consequence, intensifies the soil respira tion. However, in spite of the increase in the soil tem perature, we did not find some significant effect of the soil temperature on the rate of the soil respiration after the thinning (Table 4). The litter water content on the thinned plot significantly correlated with the soil res piration. This fact evidences that the microorganisms in the litter are highly sensitive to its moisture and play a great importance in the soil microbiological pro EURASIAN SOIL SCIENCE
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23
60 22
50 40
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20 19
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0
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(b)
697 80 70
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60 22
50 40
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Plot no.
Moisture of the soil and litter, %
Soil temperature, °C
24
Moisture of the soil and litter, % Soil temperature, °C
THE INFLUENCE OF THINNING ON THE ECOLOGICAL CONDITIONS
Plot no. 1
2
3
Fig. 2. Spatial variation of the (1) temperature and the moisture of the (2) soil and (3) litter at the (a) control and (b) thinned plots.
cesses after the thinning of the forest. Thus, although the soil respiration is greatly related to the soil temper ature and moisture, these two factors did not signifi cantly contribute to the spatial variation of the soil res piration, since their values changed little under these conditions. Another factor affecting the soil respiration rate is the larch roots. However, as our investigations showed, the root biomass was not a considerable factor affect ing the СО2 emission. No significant correlation between the soil respiration and the root biomass was found in the plots investigated (Table 4, Fig. 4). Some authors also point to the absence of relationships between the abundance of roots and the soil respira tion [12]. No significant differences were revealed between the fine root mass in the soils of the control and disturbed plots; the mean root biomass was 18% greater in the control than in the thinned plot (Table 3). This can be explained by the fact that, after thinning, 26% of the larch tree biomass and 7% of the biomass of the broadleaved trees were removed (Table 2). The den sity of the roots in the plot with the thinned stand var ied to a greater extent (CV = 88%) as compared to the control area (CV = 29%) (Fig. 4). According to Qu [22], in 2001–2003, the contribution of the root respi ration to the total СО2 emission from the soil surface in the larch forest of the National Tomakomai Forest amounted to 18 to 52%. Therefore, the 18% decrease in the root reserves after the thinning was compensated for by the increase in the microbial respiration due to the more intense decomposition of the dead roots of the cut trees. Toland and Zak [24] also reported about the activation of the microbial respiration, which compensated for the reduction of the root respiration in the first year after clear cutting. Similar data are given for broadleaved forests [17] and fir–spruce stands [18]. Thus, the fast decomposition of the dead EURASIAN SOIL SCIENCE
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roots is capable of compensating for the decrease in the root respiration in the first year after the thinning. The litter density was 15% smaller on the thinned plot than in the control (Table 3). The thinning changed not only the litter density but also affected the intensity of the soil respiration. By Qu’s results [22], in 2001–2003, the contribution of the litter respiration to the total СО2 emission in the larch forest of the National Tomakomai Forest was 8 to 32%. In addi tion, the respiration of the mineral soil layer also cor related with the litter density only on the thinned plot (Table 4). According to data of many authors, harvesfing affects the total carbon and nitrogen contents in the upper (0 to 10cm) layer of the soil. It decreases in the first year after the cutting and, then, increases during the next year [8]. Based on the analysis of the pub lished results, Johnson [14] concluded that cutting changed the carbon content in the soil by 10%. Our data agree with this conclusion, although fluctuations in the carbon content in the soil can continue during some months or even years until it reaches the initial value. We found similar dynamics of the changes in the carbon and nitrogen contents after the thinning: 7 months after the thinning, the carbon content in the upper soil layer was higher by 7% and that of nitrogen by 18% (Table 3) as compared to those in the control. Our data also agree with Qu’s results [22]: the carbon con tent in the soil under the larch forest of the Tomakomai National Forest varied from 5 to 25% and the nitrogen content from 0.2 to 1.5%. The carbon content and the C/N ratio in the soil on both plots were similar, since the nitrogen content was 18% greater in the soil of the thinned stand than in the soil of the control plot (Table 3). The carbon con tent in the 0 to 10cm soil layer varied in the control (4–23%) to a greater extent than in the soil under the
MASYAGINA et al. 20
20
18
18
Soil respiration, mmol/m2/s
Soil respiration, mmol/m2/s
698
16 14 12 10 8 6 4 18
19
20 21 Soil temperature, °C 1
22
23
16 14 12 10 8 6 4 2 0
50 100 Stock of fine roots, g/m3 1
2
150 2
Fig. 3. Relationship between the soil respiration and the temperature at the (1) control and (2) thinned plots.
Fig. 4. Relationship between the soil respiration and the density of fine larch roots in the control (1) and the thinned plot (2).
thinned stand (6–23%), whereas the nitrogen content in the same layer of the latter soil varied from 0.4 to 1.4% and, in the control, from 0.5 to 1.1%. The C to N ratio in the control soil varied more strongly (7.2– 28.4), whereas, in the soil of the thinned plot, it varied from 8.9 to 22.8, attesting to the more intense miner
alization of the falloff. The C/N values are identical to those (22–27) in the soils of pine stands (Pinus taeda L.) in Los Angeles [8]. According to Klopatek’s estimates [15], the C/N ratio was 25 in the 40yearold forest. As Gough and Seiler [12], we found a positive correlation between the soil respiration and the C/N ratio in the
Table 4. Correlation analysis of the factors studied in the control (above the lines, N = 30) and thinned (under the lines, N = 36) plots Factor
Rs
Ts
MSWC
LWC
D
LD
C
N
Rs
1 1
Ts
0.35 0.17
1 1
MSWC
0.11 – 0.04
– 0.35 – 0.32
1 1
LWC
– 0.24 – 0.41*
– 0.34 – 0.45*
0.63* 0.69*
1 1
Density of fine roots (D)
0.25 – 0.06
– 0.16 – 0.06
– 0.13 0.16
– 0.40* 0.16
1 1
Litter density (LD)
0.18 0.40*
0.27 – 0.18
– 0.36* 0.28
– 0.79* 0.09
0.27 0.15
1 1
C
0.08 0.09
– 0.18 – 0.08
0.67* 0.91*
0.37* 0.38*
– 0.13 0.01
– 0.04 0.22
1 1
N
0.06 – 0.14
0.02 – 0.22
0.42* 0.61*
0.40* 0.52*
0.02 0.21
– 0.07 0.15
0.51* 0.77*
1 1
C/N
0.10 0.39*
– 0.17 0.08
0.54* 0.56*
0.15 – 0.10
– 0.12 – 0.21
– 0.01 0.25
0.83* 0.56*
– 0.02 – 0.07
C/N
1 1
* The correlation (r) is significant at P < 0.05. EURASIAN SOIL SCIENCE
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Soil respiration, mmol/m2/s
THE INFLUENCE OF THINNING ON THE ECOLOGICAL CONDITIONS 20 18 16 14 12 10 8 6 4 2 0 500
3.
4.
5.
1000
1500 2000 Litter density, g/m3 1 2
2500 6.
Fig. 5. Relationship between the soil respiration and the litter density in the control (1) and the thinned plot (2).
7.
upper 10cm mineral layer of the soil under the larch forest after the thinning (Table 4). 8.
CONCLUSIONS The thinning of the 50yearold larch forest affected the soil temperature, the litter density, and the nitrogen content in the soil. The moisture and respira tion of this soil, as well as the carbon content, increased insignificantly. The litter moisture, the den sity of fine larch roots, and the C/N ratio decreased to some extent because of the stand’s thinning. On the whole, the thinning increased the variation of the tem perature and moisture of the soil, the density of the fine larch roots and litter, and the soil respiration. The main factor affecting the soil respiration in the larch forest is the water content of the litter.
9.
10.
11.
12.
ACKNOWLEDGMENTS The authors thank Prof. Takashi and Prof. Lai Qu from Hokkaido University for help in collecting the material. This study was supported by the Global Environmental Research Foundation of the Ministry of Environment of Japan and by the Russian Founda tion for Basic Research (project nos. 030448037 and 070496812).
14.
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EURASIAN SOIL SCIENCE
Vol. 43
No. 6
2010
