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Plant and Soil 203: 227237, 1998. 1998 Kluwer Academic Publishers. Printed in the Netherlands.

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Methane production capacities of different rice soils derived from inherent and exogenous substrates
R. Wassmann1,2,4 , H.U. Neue1,3 , C. Bueno1 , R.S. Lantin1 , M.C.R. Alberto1 , L.V. Buendia1 , K. Bronson1 , H. Papen2 and H. Rennenberg2
Rice Research Institute (IRRI), P.O. Box 933, 1099 Manila, Philippines; 2 Fraunhofer Institute for Atmospheric Environmental Research, Garmisch-Partenkirchen, Germany. 3 Present address: UfZ-Centre for Environmental Research Leipzig-Halle, Department of Soil Science, Bad Lauchstaedt, Germany. 4 Corresponding author
Received 26 January 1998. Accepted in revised form 7 July 1998
1 International

Key words: methane production, organic carbon, Philippines, rice soils, subsoil, substrate amendment, temperature effect, topsoil

Abstract Methane production rates were determined at weekly intervals during anaerobic incubation of eleven Philippine rice soils. The average production rates at 25 C varied in a large range from 0.03 to 13.6 g CH4 g(d.w.soil)1 d1 . The development of methane production rates derived from inherent substrate allowed a grouping of soils in three classes: those with instantaneous development, those with a delay of approximately two weeks, and those with a suppression of methane production of more than eight weeks. Incubation at 30 and 35 C increased production capacities of all soils, but the grouping of soils was still maintained. The Arrhenius equation provided a good t for temperature effects on methane production capacities except for those soils with suppressed production. Acetate amendment strongly enhanced methane production rates and disintegrated the grouping. However, the efciencies in converting acetate to methane differed among soils. Depending on the soil, 16.566.7% of the added acetate was utilized within ve weeks incubation at 25 C. Correlation analyses of methane production (over eight weeks) and physico-chemical soil parameters yielded signicant correlations for the concentrations of organic carbon (R 2 = 0.42) and organic nitrogen (R 2 = 0.52). Correlation indices could substantially be enhanced by using the enriched fraction of organic carbon (R 2 = 0.94) and organic nitrogen (R 2 = 0.77), i.e. the differential between topsoil and subsoil concentrations of the respective compounds. The enriched organic material in the topsoil corresponds to the biologically active fraction and thus represents a good indicator of methane production derived from inherent substrate. The best indicators of the conversion rate of acetate in different soils were pH-value (R 2 = 0.56) and organic carbon content (R 2 = 0.52). Apparently, soil properties affect methane production through various pathways. Inherent organic substrate represents a considerable source of methane in some soils and is negligible in others. Likewise, soils also differ regarding the response to exogenous substrate. Both mechanisms yield in a distinct spatial variability of methane production in rice soils.

Introduction The global methane budget has gained much attention over the last few decades. Atmospheric methane is an important factor for both the radiation balance of
FAX No: 632 891-1292. E-mail: RWassmann@irri.cgnet.com

the atmosphere and air chemistry (see GEIA, 1993; IPCC, 1990, 1992). Atmospheric methane concentrations have doubled over the last 150 years while methane release from agricultural sources is thought to be a major contributor to the increasing imbalance in global sources and sinks (Cicerone and Oremland,

228 1988). Expansion of the land used for cultivating rice and multiple copping have increased methane source strengths from main rice growing areas during the last century (Neue, 1993). Methane is generated in the last step of the anaerobic degradation chain of organic matter. The immediate precursors of methane are either acetate or CO2 /H2 . The emission of methane has been extensively investigated in eld studies (Neue et al., 1994; Sass et al., 1994; Wassmann et al., 1993), while in situ records of the two microbial processes involved, i.e. methane production and oxidation, are not available. It is generally accepted that methane production rates exceed the actual amount of methane released from the eld by factors of 24 (Holzapfel-Pschorn et al., 1985) indicating an intensive consumption of methane. Since methanotrophic bacteria can oxidize ammonia, methane oxidation is closely linked to the nitrogen cycle (Conrad and Rothfuss, 1991). The methane budget of rice elds is determined by farming practice, i.e. water management, fertilizer application etc., as well as natural factors. Previous eld experiments clearly demonstrated the site-tosite variation even under identical climate and crop management. The major objectives of this incubation study were to assess the range of methane production capacities through a survey of selected rice soils, to evaluate the response of these rice soils to exogenous substrate, to delineate relationships between physico-chemical properties and methane production, and (if possible) to identify soil properties that can be used as indicators for methane production. tems for determining methane emission measurements from rice elds (Wassmann et al., 1995). At all sites, the topsoil is generally puddled before crop establishments. Soil samples were collected during fallow period. Physico-chemical parameters of the soil samples (Table 2) were determined according to the SSSA/ASA (1996) guidelines for soil analysis: organic carbon (dichromate oxidation); organic nitrogen (Kjeldahlanalysis), phosphorus (Olsen-analysis), active Fe (extraction and AAS) as well as standard analyses for CEC, pH and texture. The enriched C- and N-contents (Table 2) were calculated by deducting the concentrations in the soil layer below the plow pan from the actual values in the soil samples. The values for the subsoil were obtained from Raymundo et al. (1989) for the soils PI (pedon #006), UR (#011), BG (#013), ML (#010), LU (#026), MH (#004), and GP (#009) and Gaunt et al. (1997) for the soils LG (soil #2), SF (#9), and LA (#3). The subsoil layer was generally identied as B-horizon; only the soils BG and LU lack a B-horizon and the respective layer was identied as AG-horizon (23-50 cm) and CR1-horizon (1933 cm), respectively (Raymundo et al., 1989). The experiments of this study addressed three different properties of methane production in soils: (1) production capacity at a given temperature, (2) production capacities at different temperatures, and (3) response of methane production to organic amendments. These approaches differed in the number of measurements required for one soil, so that the number of soils (eleven soils in Experiment A, seven in Experiment B, and nine in Experiment C) had to be adjusted to optimize measurement procedures in each experiment. Experiment A. Methane production capacities of different soils Soil preparation and incubation devices Soil samples were airdried and pulverized. All incubations were conducted with three replicates. Subsamples of 20 g pulverized soil were placed in spoutless 100 mL beakers and 40 mL distilled water was added. The beakers were closed by rubber stoppers equipped with a platinum electrode, gas inlet and gas outlet. The stoppers were sealed by silicone. A magnetic stirring bar was kept permanently inside each beaker for homogenizing the soil dispension during sampling. Incubation was started by purging the soil suspension with N2 . The gas was ushed through a glass tube

Material and methods Eleven soil samples were collected on the Philippine islands, ten at Luzon and one at Panay (Table 1). Seven of these sites were formerly integrated into the International Network on Soil Fertility and Fertilizer Evaluation for Rice (INSFER) which provided a sitespecic documentation on soil classication, chemical and physical properties as well as climatic and agronomic characterization (Raymundo et al., 1989). The soils MH (Maahas) and ML (Maligaya) represent the sites at International Rice Research Institute and Philippine Rice Research Institute, respectively, which are equipped with automated measurement sys-

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Table 1. Characterization of soils and incubation results at 25 C (methane production capacities over 2 and 8 weeks and acetate conversion rates) Soil code Location Province Soil order Methane prod. capacity (g CH4 g(d.w.soil) 1 ) 2 weeks 8 weeks 71.5 137 14.9 23.7 1.73 1.63 1.26 2.38 0.35 0.89 0.51 683 382 260 129 77.8 69.4 47.6 74.4 8.4 7.8 1.1 Acetate conversion (%) 56.4 63.8 16.5 39.7 53.5 n.r. n.r. 26.3 17.6 66.7 43.9

LG PI SD SF UR BG ML LU LA MH GP

Ligao Pila S. Dionisio S. Fernando Urdaneta Bugallon Maligaya Luisiana Lallo Maahas Gapan

Albay Laguna Iloilo Pampanga Pangasinan Pangasinan Nueva Ecija Laguna Cagayan Laguna Nueva Ecija

Mollisol Alsol Inceptisol Entisol Inceptisol Vertisol Vertisol Entisol Alsol Mollisol Inceptisol

Note: n.r. = not recorded.

Figure 1. Schematic presentation of the incubation scheme over 56 days; each interval was started by ushing with N2 ; gas sampling was conducted at the end of record phases.

Table 2. Physico-chemical properties of air-dried soil (see Table 1 for soil codes) Topsoil Soil Code pH Org. C (%) 2.35 2.08 1.2 0.71 1.58 1.62 1.21 1.41 0.88 1.86 1.18 CEC (cmol/ kg) 8.03 27.2 6.09 7.41 30.8 28.4 28.4 24.9 16.6 31.7 42.3 Org. N (%) 0.297 0.182 0.128 0.075 0.143 0.170 0.13 0.162 0.097 0.19 0.139 P (ppm) Active Fe (%) 0.44 0.756 1.23 0.361 1.11 0.649 1.15 0.388 2.23 2.27 1.04 Clay (%) Silt (%) Sand (%) Topsoil Subsoil Enr. Enr. org. C org. N (%) (%) 2.2 1.54 n.r. 0.68 0.85 0.88 0.67 0.49 0.46 0.38 0.64 0.293 0.337 n.r. 0.067 0.066 0.106 0.043 0.064 0.049 0.06 0.094

LG PI SD SF UR BG ML LU LA MH GP

5.15 7.48 3.75 5.72 6.61 6.91 6.65 4.59 5.62 6.43 6.36

4.2 24 3.5 38 18.0 3.4 3.4 5.9 4.5 28.0 18.0

8 37 25 8 31 29 59 56 23 43 54

27 34 67 22 53 57 33 40 47 44 42

65 29 8 70 16 14 8 4 30 13 4

Note: n.r. = not recorded.

230 penetrating into the soil suspension and was released via headspace through the gas outlet at the stopper. This ushing of N2 lasted for 3 min at a ow rate of 300 mL/min while the soil dispension was stirred simultaneously. Given the small volume of soil suspension (10 g soil in 40 mL water) and headspace (60 mL), this intensive ow-through of N2 removed all oxygen from liquid and gas phase in the beaker. Incubation scheme Incubation over 56 days at 25 C was divided into a sequence of phases (Figure 1). All phases started with the purging procedure as described above. Gas samples were collected at the end of each record phase, i.e. at a total incubation time of 2, 7, 14, 21, 28, 35, 42, 49 and 56 days. Connecting phases were not used for sampling, because the continuous incubation for six days led to excessively high methane concentrations in the headspace (data not shown). This incubation scheme with short record phases in weekly intervals (Figure 1) allowed a longer observation period as a previously used method with one continuous enclosure for ten days (Denier van der Gon et al., 1992). However, the pattern during the rst week was different from the ensuing weeks to allow a stabilization period. Sampling and measurement of pH/Eh Gas from the headspace was sampled with syringes and analyzed the same day. After gas sampling, a combined pH/Eh-electrode was temporarily inserted into the soil dispension through a specic access in the rubber stopper. Redox potential was determined by using the platinum electrode embedded in the stopper and a calomel reference electrode. The soil suspension was stirred throughout these measurements. Then, the gas in the headspace was discarded and the soil suspension was ushed with N2 to start a new incubation phase. Analysis Methane production rates of different soil samples were determined by analyzing a subsample of the headspace in a gas chromatograph equipped with a ame ionization detector and a Porapack N column. Methane production (P ) was calculated by P = dc V H MW T st dt W s MV (T st + T ) (g CH4 g(d.w.soil)1 d1 ) d1 ); VH the volume of headspace L), Ws the dry weight of soil (g), MW the molecular weight of methane (g), MV the molecular volume (L), T the temperature(K), and Tst the standard temperature (K). Experiment B. Methane production capacities at different temperatures Production capacities were determined at temperatures of 30 C and 35 C for seven soils (see Figure 5). This experiment followed the measurement protocol of Experiment A; results were pooled together with the ndings at 25 C. Experiment C. Response to substrate amendments In this experiment, nine different soils (see Table 2) were tested for the response to substrate amendment. For each of these soils, two sets of four replicates were initially incubated at 25 C without organic amendments for eight weeks following the procedures described above. Then, acetate (1 mL of 4.1 M acetate buffer with pH = 7) was added to one set whereas the other set was left without amendment as a control. The incubation was continued for ve weeks. Statistical analyses were done to determine the effect of soil physico-chemical properties on methane production obtained in Experiments A, B and C (Table 3). The following parameters were determined: R 2 to show the degree of association between methane production and the respective soil parameter, F -test to present the signicance of the relationship, and regression coefcient (b) to show the contribution of the independent variable being associated. Multiple regression analyses were done to determine the combined effect of several variables on methane production.

Results Methane production from inherent substrate Methane production rates of the eleven soils studied showed pronounced differences at an incubation temperature of 25 C. Temporal patterns of production rates during eight weeks of incubation (Figure 1) indicated three different classes of production patterns: class (I) instantaneous LG, PI, SD, SF class (II) delayed (for 2 weeks) UR, BG, ML, LU class (III) suppressed (for 8 weeks) LA, MH, GP

in which dc /dt is the recorded change in the mixing ratio of methane in the headspace over time (ppmv

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Table 3. Correlation analyses of physico-chemical soil properties (independent variable) and incubation results (dependent variable) Parameter pH Org. C CEC Org. N P Active Fe Clay Silt Sand Enr. Org. C Enr. Org.N

2-week capacity of methane production R2 F -test b 0.06 ns 9.9 0.34 ns 50.9 0.04 ns -0.07 0.2 ns 335.1 0.05 ns 0.7 0.12 ns -22.3 0.07 ns -0.6 0.16 ns -1.3 0.19 ns 0.8 0.59 ** 61.5 0.89 ** 404

8-week capacity of methane production R2 F-test b 0.04 ns -40 0.42 * 273.5 0.29 ns -9.7 0.52 * 2593 0.01 ns -2.4 0.2 ns -141.5 0.28 ns -6.2 0.09 ns -4.9 0.33 ns 5.2 0.94 ** 372.6 0.77 ** 1809

Acetate conversion rate R2 F -test b 0.56 * 12.6 0.52 * 25.2 0.17 ns 0.6 0.31 ns 166.4 0.3 ns 0.84 0 ns -1.42 0 ns 0.1 0.19 ns -0.6 0.04 ns 0.1 0.21 ns 12.7 0.28 ns 79.5

Note: R 2 = coefecient of determination; b = regression coefcient; * = signicant at 5% level; ** signicant at 1% level.

Figure 2. Methane production rates derived from inherent substrate of different soils (see Table 1 for soil codes); airdried soil samples were anaerobically incubated at 25 C.

232 The production rates of class (I) reached maximum values within the rst two weeks of incubation while the methane production of class (II) and (III) was still very low. The soils with a delayed development showed a steady increase of methane production rates after 3 weeks of incubation and maximum values after seven weeks. The differences between class (I) and (II) decreased within the second half of the incubation period due to decreasing production in class I and the persistently high production rates in class (II). The soils of class (III) had virtually no methane production in the rst half and only a slight increase in production rates in the second half of incubation. The results of correlation analysis between methane production rates and physico-chemical parameters are displayed in Table 3. Methane production over a 2-week incubation did not show signicant correlation to any parameter of the topsoil while methane production over eight weeks was signicantly correlated (at 5% condence interval) to organic carbon (R 2 = 0.42) and organic nitrogen content (R 2 = 0.52). Through multiple regression, the algorithm P8 = 40.7 + 37.7 OC + 2109 ON 5.4 clay could explain 72% of the differences in the soil-specific production capacities (g CH4 g(d.w.soil)1 d1 ) over an 8-week period (P8; ) using concentrations (in %) of organic carbon (OC), organic nitrogen (ON) and clay. Considerably better correlation indices were obtained by using the enriched fractions of organic carbon and organic nitrogen (Table 3). These fractions are computed by subtracting the concentrations in the subsoil from the topsoil values. Methane production rates (over two and eight weeks, respectively) were signicantly correlated to enriched carbon (R 2 = 0.59; R 2 = 0.94) as well as to enriched nitrogen (R 2 = 0.88; R 2 = 0.77). The dynamics of Eh and pH values within the rst four weeks as well as the nal values after eight weeks of incubation at 25 C are displayed in Figure 3. The soils with very low initial pH encompassed rapid changes within the rst two weeks of the incubation towards neutral pH. Development of redox potentials within the rst week revealed two distinct patterns. The group with a high production capacity was either fully reduced within 3 days or encompassed a very rapid change from high to low Eh. Eh values of the other soil classes decreased slowly from neutral to low values or from high values to neutral. The Eh and pH

Figure 3. Eh (bars) and pH (lines) at anaerobic incubation at 25 C; each subgroup comprises the sequence of records at 0, 1, 2, 3, 5 and 8 weeks for each of the soils (see Table 1 for soil codes).

values remained relatively constant during the second half of the incubation. Temperature effects on methane production Methane production rates at 25, 30 and 35 C are displayed jointly with Eh- and pH-development for one soil of each class in Figure 4. This gure also displays standard deviation for one temperature; results with other soils and temperatures showed a similar range of standard deviation. Higher temperature resulted in a faster development of the methane production rates and higher maximum values; temperature effects were especially high for soils of class II (Figure 4). However, temperature effects did not alter the classication of soils; the division in three classes was still apparent at higher temperature for the three soils shown in Figure 4 as well as for the other soils not displayed graphically. Incubation at higher temperature accelerated the reduction of the soils with low and moderate production capacity while the soil with high production capacity was already fully reduced after two days at all incubation temperatures. All three soils remained in a neutral pH range irrespective of the incubation temperature. The response of methane production to higher temperature was plotted in a logarithmic scale in Figure 5. This type of plot can be used for testing accordance with the Arrhenius equation (P = production capacity; R = gas constant; T = temperature; Ea = activation energy; a = Arrhenius constant): P = a eEa /RT A soil responding to temperature changes according to the Arrhenius equation is reected by a linear

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Figure 4. Methane production capacities, Eh and pH during anaerobic incubation of the soils MH, LU and PI at 25, 30 and 35 C (see Table 1 for soil codes); standard deviations are only depicted for one temperature (at 25 C) to prevent graphic overlaps.

graph in this type of plot. The seven soils tested in this experiment are represented by sets of three data points. The graphs of the soils with a high and moderate production were roughly linear (R 2 > 0.8) indicating accordance with Arrhenius equation. For the soils of group (III), however, the production rates at 25 and 30 C were virtually zero and the temperature response was not linear (R 2 < 0.8). Activation energies ranged from 47 to 114 kJ mol1 K1 but did not show a relationship to soil classes. Methane production from exogenous substrate The addition of acetate greatly enhanced methane production rates of all soil types and completely disintegrated the classication according to inherent production capacities (Figure 6). The production rates triggered by acetate amendments varied for different soils from average values of 25 to 65 g CH4 g(d.w.) 1

d1 . A stochiometric assessment of the methane produced in different samples yielded that 16.5%66.7% of the acetate supply was utilized in the course of this incubation period (Table 1). Correlation analysis of physico-chemical soil properties indicated a signicant correlation with soil pH (R 2 = 0.56) and organic carbon (R 2 = 0.52) while the R 2 -value obtained by multiple regression with both parameters was only marginally higher (R 2 = 0.57).

Discussion The airdrying of the samples roughly corresponds to the typical soil treatment in the eld during the fallow period. Different inoculation of samples due to soil drying can largely be excluded because the density of methanogens in rice soils is not markedly decreased

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8
ln CH4 production [g CH 4 g(soil d.w.)-1]

7 6 5 4 3 2 1 0 3.20

GP MH LU ML BG UR PI

3.25

3.30 1/T (10-3 * K-1)

3.35

3.40

Figure 5. Arrhenius diagram of methane production capacities at 25, 30 and 35 C (corresponding to 3.36, 3.3 and 3.25 * 10-3 K1 , respectively).

by the dry fallow period (Joulian et al., 1996). Furthermore, the initiation of methanogenesis is generally not limited by number of methanogens (Mayer and Conrad, 1990). The eleven soils showed a pronounced variation in methane production capacity which is in line with previous ndings. A survey of 16 soils from USA, India, Thailand and Liberia also documented that production capacities varied over several orders of magnitude (Wang et al., 1993b). Likewise, a preliminary screening of 20 Philippine soils of which eight were also integrated in the study presented here showed significant differences in methane production after ten days of incubation (Denier van der Gon et al., 1992; Neue et al., 1994). In this new study presented here, soils were classied according to development patterns (instantaneous, delayed, and suppressed) which appeared to be a more reliable criterion than the magnitude of production. The temporal patterns observed in this study are in line with ndings by Wang et al. (1993b) whereas classications according to total production were highly dependent on the duration of the observation period. The classication according to temporal development was not affected by different incubation temperatures. The increment in methane production due

to higher temperature was considerably lower than the inherent differences between the three soil classes (Figures 4 and 5) so that the grouping persisted throughout the temperature range from 2535 C. The temperature dependency of methane production rates was shown for several soils individually (Conrad et al., 1987; Sass et al., 1991; Schtz et al., 1990). This study corroborated these ndings for a larger number of soils with different production capacities (Figure 5). The Arrhenius equation generally provided a good t for the temperature effect on methane production as previously shown by Sass et al. (1991). Only in the soils with suppressed production this temperature effect was not detectable. In this study, methane production capacity from inherent substrate was signicantly correlated to the organic carbon and organic nitrogen content. The organic fraction in the soil consists of an array of different compounds including methanogenic material. However, the retarded development of methane production of group (II) resulted in an insignicant relationship for the 2-week incubation, and even the correlation indices for 8-week incubation were not high enough to postulate a robust pedo-transfer function. This is in line with other studies. Wang et al. (1993b) concluded that no individual chemi-

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70 60 gCH4 g(soil d.w.) -1 d-1 50 40 30 20 10 0 SD PI LG

Acetate (9-13 weeks) No acetate (9-13 weeks) Initial (0-8 weeks)


GP

SF UR LU LA MH

Soil type

Figure 6. Average methane production rates derived from inherent substrate (08 weeks) and continuing incubation (913 weeks) without and with acetate amendment at 25 C (see Table 1 for soil codes).

cal parameter could be singled out as an indicator for methane production capacity. Multiple regression analysis yielded a substantially higher correlation index (R 2 = 0.72) combining organic nitrogen, organic carbon, and clay content in one equation. The role of clay in methane production can be explained by a protection of organic matter from microbial breakdown by clay particles (Oades, 1988). The enriched fractions of organic carbon and organic nitrogen, i.e. the difference between the puddled layer and the layer below the plow pan, proved to be a very good indicator for methane production (Table 3). This nding can be explained by the hypothesis that the differential in organic matter between topsoil and subsoil corresponds to the biologically active fraction in the soil (Gaunt et al., 1997). Based on this assumption, high contents of enriched organic material will ultimately translate into high methane production in the soil. However, data obtained in previous incubation experiments conducted at IRRI (Denier van der Gon et al., 1992) did not show signicant correlations to

enriched carbon (Gaunt et al., 1993). The newly introduced incubation scheme (Figure 1) allowed much longer incubation periods and implied low standard deviations which can be regarded as decisive factors for proving statistically signicant relationships. The close correlation to enriched nitrogen will not differ substantially if total nitrogen is used in stead of organic nitrogen because the organic fraction comprises more than 90% of the total nitrogen in the soil (SSSA/ASA, 1996). Methane production can be impeded by low and high pH and by a high redox potential (Ponnamperuma, 1972; Wang et al., 1993a). The initial pH of the soils of this study varied broadly. However, incubation rapidly neutralized pH in all soils (Figure 3) which has also been found by Ponnamperuma (1972). Therefore, pH can be excluded to have a direct impact on the development of methane emission in the latter stages of the incubation. The redox potential reects chemical and biochemical processes that depend on the nature of available organic matter and the buffering capacity of the soil (Neue, 1988). Thus, the dynamic changes

236 in soils with high production rates (Figure 3) can be attributed to intense bacterial degradation of organic material exceeding the availability of oxidants. The inherent production capacity is apparently determined by an interaction of various chemical and physical parameters under anaerobic conditions. Acetate is one of the main precursors of methane production in rice soils (Schtz et al., 1989; Takai, 1970) and can either derive from root exudation (Lin and You, 1989) or from fermentation (Neue and Roger, 1993). The conversion rates of acetate to methane differed greatly among soil types and were signicantly correlated to organic carbon and pH. However, acetate was added after eight weeks of incubation when the samples were already depleted in the organic content and were in neutral pH range. More data on the dynamics of soil parameters during incubation are needed to explain different response patterns of methane production to exogenous amendments. References
Cicerone R J and Oremland R S 1988 Biogeochemical aspects of atmospheric methane. Global Biogeochem. Cycles 2, 299327. Conrad R and Rothfuss F 1991 Methane oxidation in the soil surface layer of a ooded rice eld and the effect of ammonium. Biol. Fertil. Soils 12, 2832. Conrad R, Schtz H and Babbel M 1987 Temperature limitation of hydrogen turnover and methanogenesis in anoxic paddy soil. FEMS Microbiol. Ecol. 45, 281289. Denier van der Gon H A C, Neue H U, Lantin R S, Wassmann R, Alberto M C R, Aduna J B and Tan M J 1992 Controlling factors of methane emission from riceelds. In World Inventory of Soil Emission Potentials. Eds. N H Batjes and E M Bridges. pp 81 92. WISE Rep. 2, ISRIC, Wageningen. Denier van der Gon H 1996 Methane emission from wetland rice elds. Ph.D. Thesis, Agric. Univ. Wageningen, The Netherlands. 182 p. Gaunt J L, Neue H U, Bragais J, Grant I F and Giller K E 1997 Soil characteristics which regulate soil reduction and methane production in wetland rice soils. Soil Sci. Soc. Am. J. (In press). GEIA Global Emission Inventory Activity 1993 Report on the 3rd workshop (Amersford 31 Jan02 Feb. 1993). Ed. A F Bowman (RIVM-Rep. 481507002, Bilthoven, The Netherlands. Holzapfel-Pschorn A, Conrad R and Seiler W 1985 Production, oxidation and emission of methane from rice paddies. FEMS Microbiol. Ecol. 31, 343351. IPCC Intergovernmental Panel on Climate Change 1990 Climate Change. The IPCC Scientic Assessment. Cambridge University Press. XXXIX + 358 p. IPCC Intergovernmental Panel on Climate Change 1992 Climate Change. The Supplementary Report to the IPCC Scientic Assessment. Cambridge University Press. XII + 200 p. Joulian C, Ollivier B, Neue H U and Roger P A 1996 Microbiological aspects of methane emission by a riceeld soil from the Camargue (France): 1. Methanogenesis and related microora. Eur. J. Soil Biol. 32(2), 6170. Lin M and You C 1989 Root exudates of rice (Oryza sativa L.) and their interaction with Alcaligenes faecalis. Sci. Agric. Sin. 22, 612. Mayer H P and Conrad R 1990 Factors inuencing the population of methanogenic bacteria and the initiation of methane production upon ooding of paddy soil. FEMS Microbiol. Ecol. 73, 103 112. Neue H U 1988 A holistic view of chemistry of ooded soils. In IBSRAM International Board of Soil Research Management. Soil Management for Sustainable Rice Production in the Tropics. pp 332. Monogr. No. 2, Bangkok, Thailand. Neue H U 1993 Methane emission from rice elds. Bioscience 43, 466474. Neue H U and Roger P A 1993 Rice agriculture: Factors controlling emissions. In Global Atmospheric Methane. Ed. M A K Khalil and M Shearer. pp. 254298. NATO ASI/ARW series. Neue H U, Lantin R L, Wassmann R, Aduna J B, Alberto M C R and Andales M J 1994 Methane emissions from rice soils of the Philippines. In CH4 and N2 O: Global Emissions and Controls from Rice Fields and Other Agricultural and Industrial Sources. Ed. K Minami, A Mosier and R Sass. pp 5563. NIAES Series 2, Tsukuba, Japan. Oades J M 1988 The retention of organic matter in soils. Biogeochemistry 5, 3570. Ponnamperuma F N 1972 The chemistry of submerged soils. Adv. Agron. 24, 2996.

Conclusions Apparently, soil properties affect methane production through various pathways. Inherent organic substrate represents a considerable source of methane some soils and is negligible in others. Superimposed on this variation regarding inherent substrate are different responses to exogenous substrate that can be derived from different conversion rates of acetate. The interaction of both properties may explain spatial and temporal variability of methane production, and in the next level methane emission of rice elds. Methane production capacities showed good correlations to the enriched organic fraction of soils. These data can be derived from advanced soil data bases. Therefore, the results of this study open up the opportunity to integrate soil information into site-specic assessments of methane production and emission from rice elds. However, the scope of this study with a limited number of rice soils demands for more comprehensive surveys on methane production in major soils of different rice growing areas.

Acknowledgement This work was conducted within the project An Interregional Research Programme on Methane Emissions in Riceelds funded by UNDP/GEF (GLO/91/G31).

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