Agroforestry Systems 57: 117125, 2003. 2003 Kluwer Academic Publishers. Printed in the Netherlands.

117

Characteristics and decomposition rates of pruning residues from a shaded coffee system in Southeastern Brazil
E.S. Mendona 1,2,* and D.E. Stott 1
USDA Agricultural Research Service, National Soil Erosion Research Laboratory, IN 47907-1196 West Lafayette, USA; 2Departamento de Solos, Universidade Federal de Viosa, 36570-001 Viosa, Minas Gerais, Brazil; *Author for correspondence (e-mail: esm@mail.ufv.br fax: 31-38992648)
Received 8 April 2002; accepted in revised form 30 December 2002
1

Key words: Century simulation, Lignin, Nitrogen, Shaded perennial system, Tropical trees Abstract In the Zona da Mata Mineira of Southeastern Brazil the development of sustainable land requires the integration of crops with trees. The objectives of this study then were to (i) characterize prunings from the main tree species in an agroforestry system; (ii) determine the effects of the physical and chemical characteristics of the prunings on their decomposition patterns in the laboratory; (iii) assess the effect of mixing leaves of different species on decomposition rates; and (iv) propose a decomposition index for the residues studied. The study was carried out with pruning residues from Cajanus cajan, Solanum variable, Cassia ferruginea, Piptadenia gonoacantha, Croton urucurana, and Melinis multiora. The materials were characterized for total C, N, P, Ca, Mg and K contents; lignin, cellulose, hemicellulose and soluble polyphenols contents. The pruning residues had high polyphenols and lignin contents, high C:N and C:P ratios, and low contents of Ca, Mg, and K. The low decomposition rates of the prunings were related to the P, K, hemicellulose and polyphenol contents. The rates of N mineralization from most of the residues indicate that there is a potential to supply the needs of a crop of maize. The residues of some species, if decomposed alone, would not supply sufficient nutrients, and need to be mixed with leaves of other species. Introduction The Zona da Mata Mineira in Southeastern Brazil is characterized by a hilly, rolling landscape. The soils are highly weathered and acidic, with low natural fertility. At present, its agricultural sector is characterized by the use of traditional non-mechanized management systems with little or no additions of fertilizer and low food production. Under these conditions, the development of sustainable land use seems to demand erosion control and an integration of crops to increase soil biology and chemical characteristics (Franco et al. 1994). Hence, agroforestry systems may present the potential to solve part of the regions agricultural problems of the farmers since it can contribute to reduce soil erosion and increase nutrients and carbon cycle. The aims of agroforestry systems in highland areas are to increase physical protection against soil erosion and improve soil fertility. With crops, such as coffee, agroforestry may improve soil structure and increase soil organic matter content and plant-available nutrients (Palm 1995; Sanchez 1995). The success of an agroforestry system is related to the amount and quality of the tree prunings, the amount of nutrients released from the prunings during the decomposition process, and how the amount and timing of the released nutrients satisfy the needs of the crop. Several attempts have been made to quantify plant residue quality and its relation to the decomposition process (Palm and Sanchez 1991; Constantinides and Fownes 1994a; Tian et al. 1995; Handayanto et al. 1995). Residue quality is usually dened in relation to its chemical composition. The C, N, P, lignin, and polyphenolic contents, along with their interrelations,

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Table 1. Chemical characteristics of the top 15-cm soil (Typic Haplustoxs) 1 of the shaded coffee system used in the study from southeastern Brazil Soil Clay (%) Sand (%) Organic C (g kg 1) Total N (g kg 1) pH Effective Aluminum CEC Saturation (cmol kg 1) (%) 2.14 3.74 40 13 Total CEC, Available P at pH 7 (mg kg 1) (cmol kg 1) 8.5 7.6 0.8 0.9

Paulo-Vicente Farm Joo dos Santos Farm Incubation Soil

1

45 39

46 50

32 21

1.6 1.5

4.6 5.4

37

55

34

3.7

5.3

4.15

30

13.1

0.8

Latossolo Vermelho-Amarelo (Red-Yellow Latosols) in the Brazilian taxonomic system.

are the most common measurements (Sinsabaugh and Moorhead 1994; Vanlauwe et al. 1997). Residues with high decomposition and N-mineralization rates are usually considered to be of high quality. However, if the main aim of an agroforestry system is to keep the soil surface covered for erosion control, the concept of low quality residues should be reconsidered. The focus of the residue quality measurements should be on determining the dynamics of the decomposition process and nutrient cycling in different ecosystems and relating the quality with the goals of each situation. Palm (1995) pointed out that agroforestry species with high N, lignin, and polyphenolic contents may provide nutrient release patterns that more closely match the demands of crops for nutrients, especially N. Despite increasing experience with agroforestry systems in the tropics, there is still a lack of information about the characteristics and decomposition dynamics of prunings from tropical trees and bushes in our region. Most published studies dealing with residue decomposition in agroforestry systems consider only the leaves, and ignore other parts of the plant. Therefore, the objectives of this study were to: (i) characterize the prunings from the main plants of an agroforestry system in southeastern Brazil; (ii) determine the effect of the characteristics of the pruning residues on their decomposition patterns as measured in a laboratory incubation study; (iii) assess the effect of mixing leaves from different species on the decomposition rate since in the eld the plants are in the same area and the mixture of residues may change the decomposition pattern; and (iv) propose a decomposition index for the residues studied.

Material and methods Pruning residues Pruning residues were obtained from two farms in April 1997. The farms used agroforestry systems, with coffee as the main crop. The farms of Paulo-Vicente and Joo dos Santos, were located within 2 km of one another, at 203959 S latitude, 423115 W longitude, and 900 m altitude. The main soil on the farms was classied as a Latossolo Vermelho-Amarelo (Red-Yellow Latosol) in the Brazilian taxonomic system and as a Typic Haplustox in the U.S. soil taxonomic system (Table 1). Pruning residues from four tree species were collected from both sites: Cajanus cajan (common name: guandu) of the Caesalpinoideae, Solanum variable (capoeira branca) of the Solanaceae, Cassia ferruginea (cana stula) of the Caesalpinoideae, and Piptadenia gonoacantha (jacare) of the Mimosoideae. Pruning residues from two other species were also collected at the Santos farm: Croton urucurana (adrago) of the Caesalpinoideae, and a tall grass, Melinis multiora (capim gordura) of the Gramineae. The residues were divided into their component parts: leaves, the stems attached to the leaves (petioles), and the small stems (diameter <2 cm), and airdried and stored until use at ambient temperature. Sub-samples of each type of pruning component was ground with a mill and pestle to <1 mm and ashed in a muffle furnace at 450 C for 4 h. Total C and N content were analyzed by dry combustion (LECO600 CHN, St. Joseph, MI, USA). Total P was determined by the molybdenum blue method after digestion with perchloric and nitric acids (Anderson and Ingram 1989), and on the same acid digestion solu-

119 tion, Ca and Mg contents were determined using an atomic absorption spectrophotometer (GBC, Model 908AA, Victoria, Australia) and K content was determined using a ame spectrophotometer. Water potential of the residues at given water contents were measured with a thermocouple psychrometer (Tru Psi model SC10X, Decagon, Inc., Pullman, WA) (Myrold et al. 1981). To measure the water potential of the residues at various water contents, airdried leaves, petioles and stems were cut in 0.50 to 1.0-cm lengths. Samples were wetted with double deionized water to contents of 100, 200, 300, 400, 500, and 600%, by weight. The samples were allowed to equilibrate in closed plastic vials at room temperature for 24 h before measurement. Lignin, cellulose and hemicellulose content were determined by the acid-detergent ber method (Goering and Van Soest 1970). Sub-samples were placed in hot (80 C) 50% aqueous methanol for 1 h to extract soluble polyphenols. A 2 mg mL 1 plant tissue: extractant ratio was used (Constantinides and Fownes 1994b). The nal concentrations were determined colorimetrically using the Folin-Denis reagent with tannic acid as a standard (Anderson and Ingram 1989). Carbohydrates were determined by the acid hydrolysis method. All residue data are reported on an oven dry, ash-free basis. Incubation procedure The 120-d incubation experiment was conducted to determine the amount of CO 2 evolved and mass lost from the pruning residues. Pruning residues were chopped into 4 to 5 cm long pieces. Each of six treatments consisted of the leaves, petioles, and stems from a single species mixed in the same ratio as the original, uncut sample. Three additional treatments consisted of mixtures of the six types of leaves. Mixture 1 contained equal portions of leaves from C. cajan, S. variable, C. fernuginea, and P. gonoacantha; Mixture 2 included leaves, in equal portions, from C. cajan, S. variable, C. fernuginea, P. gonoacantha, and C. urucurana; and Mixture 3 contained equal amounts of leaves from C. cajan, S. variable, C. fernuginea, P. gonoacantha, and M. multiora. The soil used for the incubation experiment was a Typic Haplustox collected from the 0 to 15 cm depth (Table 1). The soil was sieved (<2 mm) to remove roots and organic debris. One hundred grams of soil was placed into each incubation jar, and 5 g of dry residue, in the appropriate ratio, were spread evenly on the soil surface (about 34 cm 2 area) within an incubation jar. One-pint Ball mason canning jars (480 ml capacity) with self-sealing lids were used as incubation jars. Double deionized water was added in sufcient quantity to bring the water content of the soil and residue to 0.33 MPa. The incubation jars, which included a soil-only control, were arranged in a completely randomized design, with six replications, in an incubator set at 25 2 C. Inside each jar was a 10 ml beaker containing 3 ml of 3 M NaOH, which was used to trap the respired CO 2. Periodically, the jars were removed from the incubator, the NaOH was replaced with fresh solution, sufficient water was added to bring the soil and residue back to the original water content, and then the resealed jars were returned to the incubator. The amount of respired CO 2 trapped in the NaOH that was removed was measured potentiometrically (Golterman and Clymo 1969). At the end of incubation period, the mass remained were separated from the soil, washed with deionized waster and measured. Incubation simulation with CENTURY Estimates of CO 2 respiration and pruning residue decomposition rates were made using the CENTURY model, Agroecosystem Version 4.0 (Metherell et al. 1993). The simulations were done with the Soil Organic Matter submodel, using Microcosms, the soil incubation mode of the model. The litter level, lignin content of the litter, and the C:N, C:P, and C:S ratios of the litter were specied from the experimental data. The constant decomposition rate coefficient (k) was estimated from the mass loss from the prunings as simulated by CENTURY, using a single exponential equation, y = X 0 exp (kt); where y is the percentage of mass remaining at time t, X 0 is the initial (t = 0) amount of mass, and k is the decomposition rate constant. Statistical analysis The differences between pruning residue characteristics were tested by analysis of variance (ANOVA). The mean values were compared using the Tukey multiple comparison method. Statistical analyses were done with SAEG, a statistical package developed by the Federal University of Viosa, Minas Gerais State, Brazil.

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Table 2. Initial chemical and physical characteristics of the tree- and bush-pruning residues from the shaded coffee system in southeastern Brazil Plant species Plant part Relative proportion a C N P Ca Mg K LG b CL c HC d CR e PP f WC h g g 1 121 191 215 176 189 207 137 180 161 156 165 216 146 161 168 307
d

------------ g kg 1 -----------C. cajan Leaf Petiole Stem Leaf Petiole Stem Leaf Petiole Stem Leaf Petiole Stem Leaf Petiole Stem Leaf 0.45 0.35 0.20 0.33 0.37 0.30 0.38 0.34 0.28 0.35 0.40 0.42 0.29 0.29 0.42 1.00 540 510 510 500 490 510 550 520 530 510 510 520 520 480 510 490 47 16 13 39 24 13 26 11 09 38 17 14 35 15 12 19 2.4 0.7 0.6 1.2 0.8 0.2 1.2 0.4 0.3 1.3 0.7 0.5 1.0 0.4 0.4 1.0 0.08 0.04 0.02 0.12 0.08 0.05 0.06 0.05 0.04 0.09 0.05 0.04 0.10 0.14 0.03 0.03 0.02 0.01 0.01 0.03 0.03 0.01 0.03 0.02 0.01 0.03 0.01 0.01 0.03 0.03 0.01 0.02
b

0.10 0.07 0.05 0.12 0.15 0.06 0.06 0.06 0.03 0.09 0.08 0.05 0.06 0.07 0.03 0.11

111 124 129 75 96 146 90 139 157 106 138 147 84 98 203 74
c

194 418 466 224 335 466 168 473 496 181 409 473 150 275 412 304

406 755 838 521 644 829 430 802 790 426 706 840 401 586 738 764

88 12 09 37 13 08 193 82 33 151 14 10 189 131 33 35

8.22 11.03 22.45 6.04 7.23 12.28 8.31 10.12 36.26 8.73 8.07 7.41 12.80 18.94 27.90 5.54

S. variable

C. fernuginea

P. gonoacantha

C. urucurana

M. multiora
a e

The proportion of the harvested residue of a plant species that is leaf, petiole, or stem. LG = lignin; CL = Cellulose; HC = Hemicellulose; CR = Carbohydrates; fPP = Polyphenols; gWC = Water Content of the residues at 0.33 MPa. The standard deviation for C, N, P, Ca, Mg, and K ranged from 5 to 10%, and ranged from 5 to 15% for all other analyses.

Results and discussion Characteristics of pruning residues Leaves, petioles, and stems varied in their chemical composition (Table 2). The elemental concentrations were low and followed the general trend of leaves > petioles > stems. The amount of N in the leaves ranged from 26 (C. fernuginea) to 47 g kg 1 (C. cajan) in the leaves and from 11 (C. fernuginea) to 24 g kg 1 (S variable) in the petioles. The M. multiora blades had lower N concentrations than the tree leaves, 19 g kg 1. The N in the stems was about the same for all species, ranging from 9 to 13 g kg 1. The P contents of the tree leaves of all species varied from 1 to 2.4 g kg 1. Among the trees, S. variable had the highest K (120 mg kg 1) and Ca (120 mg kg 1) contents in the leaves. The M. multiora blades had lower Ca concentrations when compared to the tree leaves, and the K concentration was similar to the highest K concentrations in the tree leaves. The Mg content was about the same in all the leaves, petioles, and stems.

The concentration of N and P in the leaves (Table 2) of the pruning residues, assuming an input of 10 t ha 1 yr 1 of dry matter what is the dry matter normally produced in the areas, is potentially enough to meet the amount of nutrients required to produce 2 t of maize (Palm 1995). The residues with the lower concentrations of Ca, Mg or K, would not supply sufficient amounts of these nutrients to meet the nutrient demands for the next maize crop. Leaves generally had lower contents of lignin, cellulose and hemicellulose and higher amounts of polyphenols than the petioles and stems (Table 2). Petioles showed the same trend when compared with stems. Amongst all the species, lignin contents ranged from 74 to 111, 96 to 139, and 129 to 203 g kg 1 in the leaves, petioles, and stems, respectively. Cellulose concentrations ranged from 150 to 304, 275 to 473, and 412 to 496 g kg 1 in the leaves, petioles, and stems, respectively. Contents of hemicellulose in the leaves varied from 121 to 307 g kg 1, while contents in the petioles and stems ranged from 161 to 191 and from 161 to 216 g kg 1, respectively. Total carbohy-

121 mineralized and loss of mass were low and were inuenced by the species of plant (Table 3). Among the trees, C. fernuginea produced the lowest and S. variable the highest cumulative C evolved as CO 2 and loss of mass, respectively, after 120-d of incubation. Mixing leaves of different species together signicantly increased the total C evolved as CO 2, when compared to single species residues of C. fernuginea, and decreased when compared to S. variable, C. cajan, or M. multiora. For ve residue samples, the N concentration in the residues increased during the incubation, while the concentration decreased in 3 samples and remained unchanged in one leaf mixture (Table 3). The total N in the C. fernuginea residue was low, less than 17 g kg 1 when the leaves, petioles and stems were combined in normal ratios, so if this were the only residue present, net N immobilization would occur for relatively long periods. The total N present in the residues of the other plant species was sufficient to result in net mineralization. Thirty-nine percent or less of the N in the residues was mineralized during 120 days of incubation (Table 3). The amount of N mineralized was calculated from the initial and nal N concentrations and residue mass. From 2.5 to 39% of the N was mineralized during the 120-d incubation, with S. variable residues showing the highest N mineralization potential and C. fernuginea the lowest. Several laboratory decomposition studies using leaves from leguminous trees recovered less than 50% of the initial N content (Palm and Sanchez 1991; Tian et al. 1992; Constantinides and Fownes 1994a). The net N mineralization of the pruning residues was correlated with N to (lignin + polyphenol) ratio, with r = 0.77 (P < 0.01), this ratio, developed by Palm and Sanchez (1991), increases as the total net mineralization decreases. Comparing the average daily rate of CO 2 evolved as C during the rst 21-d and the initial elemental and chemical concentrations of the residue (Table 4), there were statistically signicant correlations with P (r = 0.92), K (0.61), hemicellulose (0.63), polyphenols (0.76), and carbohydrates (0.56). When correlated with the rate of C-CO 2 evolved per day, averaged over the 120-d incubation, or with the total cumulative amount of C evolved as CO 2, P (r = 0.87), K (0.70), hemicellulose (0.63), and polyphenol (0.68) contents were signicantly related. Initial elemental and chemical concentrations that were significantly related to the mass remaining at the end of

Figure 1. Water potential curves for the different components of the Cajanus cajan pruning residues from the shaded coffee system in southeastern Brazil.

drate concentrations varied from 401 to 764, 586 to 802, and 738 to 840 g kg 1 in the leaves, petioles, and stems, respectively. Concentrations of polyphenolics in the plant material varied from 35 to 193, 12 to 131, and 8 to 33 g kg 1 in the leaves, petioles, and stems, respectively. The polyphenolic contents of the leaves were in the same range as that found by Constantinides and Fownes (1994b) for tropical plant leaves. All the pruning residues displayed similar water sorption patterns (Figure 1). At the same water contents, the stems had the highest water potential, followed by petioles and leaves. Among the trees, S. variable presented the lowest and C. urucurana the highest water potential at a given water content for the leaves and petioles. For the thick stem components, P. gonoacantha presented the lowest and C. fernuginea the highest values. This information was used to determine the water content necessary to bring the residues to a water potential of 0.33 MPa for the incubation experiments (Table 2), which will have inuence on the biomass activity that decompose the residues. Laboratory Incubations During the rst 21-d of the 120-d incubation experiment, C evolved as CO 2 at exponential rates from all residue samples. After 21 days, the rate of CO 2 evolution proceeded linearly. The total amounts of C

122
Table 3. C and N concentration in the tree- and bush-pruning residues from the shaded coffee system in southeastern Brazil before and after the 120-d incubation at 25 C, and the total amount of N mineralized during the residues incubation Plant species C content of residue Initial N content of residue Mineralized N b N Lost Portion of Initial N % 30.0 39.4 2.5 3.5 13.2 20.4 17.3 24.7 25.2

Final Initial Final ------------ g kg 1residue -----------20 a 11 30 22 11 12 32 13 22 500 470 490 470 480 450 500 480 490 11 24 13 11 32 24 11 12 32 29 24 16 23 19 19 37 36 33 2 3 1 1 2 1 4 2 1 27 20 19 29 22 21 41 35 33 1 2 1 2 1 2 4 3 1

C. cajan S. variable C. fernuginea P. gonoacantha C. urucurana M. multiora Mixture 1 Mixture 2 Mixture 3

520 500 530 520 500 490 520 520 520

8.8 9.5 0.4 0.8 2.5 3.9 6.4 8.9 8.3

a Numbers represent the mean value with standard deviation, and are based on the contents of the leaves, petioles and stems in the ratios found at harvest. bCalculated from the initial and remaining N contents of the residue.

Table 4. Correlation coefficient (r) between the initial chemical composition of the tree- and bush-pruning residues from the shaded coffee system in southeastern Brazil, with leaves, petioles, and stems in the ratios found at harvest, and the average rate of C evolved as CO 2 during the rst 21 days of decomposition, the average rate of C evolved as CO 2 during the entire 120-day incubation, and the cumulative amount of C lost as CO 2 during the 120 day incubation Average Rate of C-CO 2 Evolution During the First 21-d of incubation Full 120-d incubation a ------------ r -----------0.57* 0.16 0.92** 0.61* 0.22 0.22 0.37 0.06 0.63** 0.76** 0.56* 0.12 0.90** 0.02 0.68** 0.57* 0.46 0.53* 0.02 0.87** 0.70** 0.19 0.14 0.47 0.09 0.63** 0.68** 0.44 0.28 0.88** 0.21 0.60* 0.67** 0.61*

Initial chemical characteristics

Mass remaining after 120-d

C N P K Ca Mg Lignin (LG) Cellulose (CL) Hemicellulose (HC) Polyphenols (PP) Carbohydrates (CR) C:N C:P N:LG N:PP N/(LG+PP) (C:N)*LG/CR0.05

0.48 0.19 0.82** 0.72** 0.02 0.01 0.45 0.20 0.49 0.58* 0.33 0.44 0.89** 0.33 0.58* 0.72** 0.70**

*,** Signicant at P < 0.05 and P < 0.01, respectively. aCorrelations are the same as for the total cumulative C evolved as CO 2.

120-d included P (r = 0.82), K (0.72), and polyphenols (0.58). The concentrations of hemicellulose in the residues used for the incubations ranged from 147 g kg 1 in leaf mixture 2 to 307 g kg 1 in M. multiora. For carbohydrates, the concentrations range from 431

g kg 1 in leaf mixture 2 to 695 g kg 1 in P. gonoacantha. Polyphenol contents varied from 19 g kg 1 in S. variable to 132 g kg 1 in leaf mixture 2. There is some evidence in the literature that C, after the relatively available C is utilized, becomes limiting the microorganisms activity (Stott et al. 1989). Diack

123 (1994) observed a strong correlation between hemicellulose and carbohydrate contents and the degradation of a wide variety of residues from the temperate zone. The negative correlation of polyphenol concentrations and their associated rates of degradation have been noted by Palm and Sanchez (1991). Correlations between the decomposition rates at 21 and 120-d, as well as the mass of residue remaining after 120-d, and the residue quality indices found in the literature (Palm and Sanchez 1991; Constantinides and Fownes 1994a; Handayanto et al. 1995; Gijsman et al. 1997) (Table 4) were mixed. The N content of the residues was not correlated with the decomposition rates in this study, the lignin concentration was only weakly correlated, and the C to N and N to lignin indices did not correlated well. The N to polyphenol and N to (lignin + polyphenol) indices were moderately correlated, but the correlations were not nearly as strong as polyphenol alone. The [C to N*lignin]/ carbohydrate 1/2 index was also moderately correlated with the rates of decay, but there was a strong correlation with mass loss (r = 0.70), basically due to the weak C to N correlation being negated by the weak lignin correlation. The C to P ratio was high, ranging from 931(C. fernuginea) to 321 (M. multiora), reecting the low P availability in the strongly P-sorbing Latosols (Oxisols) of the region. The high C:P ratio in the residues, along with the somewhat high C to P ratio (43) of the incubation soil, likely resulted in the immobilization of P by the soil microbial community during the initial stages of decomposition (Ofori-Frimpong and Rowell 1999). Along with the initial P concentrations, the C to P index, correlated well with the decomposition rates (r = 0.90, r = 0.88) and mass remaining (r = 0.89), indicating that P was a controlling nutrient within the incubation system. P availability has a strong impact on litter decomposition in tropical, Pdecient soils. In these soils, residues decayed slowly, with soil microbes expending relatively more metabolic energy to acquire the decient nutrient rather than to produce the lignocellulases necessary to decay the residues to obtain C (Sinsabaugh and Moorhead 1994). The C to N ratio, ranging from 14.0 for the Leaf Mixture 1 to 33.1 for the C. fernuginea, was poorly correlated with the decomposition rates (r = 0.12, 0.28) and the amount of mass remaining at the end of 120-d (r = 0.44). There was, probably, enough N, or nearly so, to supply the needs of the microbial population involved in the decay process. This is in some conict with the studies of Vitousek et al. (1994) and Gijsman et al. (1997), however, C to N ratios of the residues in those studies were somewhat higher ranging from, 98 to 130, and 24 to 137, respectively. CENTURY model simulation The initial chemical composition and mass data for the pruning residues and the incubation conditions were used as input for the CENTURY model. For the rst run of CENTURY using these data, and the default value of 14.8 for the maximum surface metabolic decomposition rate, the model overestimated the amount of decomposition after 120 days (Table 5). This overestimation was most likely to the low nutrient status and high polyphenol content of the pruning residue, due to of the inclusion of petioles and small stems. The default value was therefore adjusted to 8.8, resulting in a simulated rate of C-CO 2 evolution similar to the experimental data. CENTURY works with lignin:N and C:N ratios to determine the amount of plant parts that go to the metabolic and structural compartments. However, some works have pointed out the importance of polyphenols content on the initial stage of plant decomposition (Palm and Sanchez 1991; Constantinides and Fownes 1994a), and the inuence of P content and others nutrients on the plant decomposition rate (Vitousek et al. 1994). Therefore, our data suggest that some adjustments, i.e. including polyphenols and P contents, should be made in the CENTURY when working with tropical plant since the decomposition rate of tropical pruning species is mainly inuenced by the (lignin+polyphenol):N ratio and P content.

Conclusion Our study shows that the plants used in the tropical agroforest presents very high polyphenols and low P contents, indicating that they are the controlling compounds within the incubation system and inducing low decomposition rates of the prunings. The species used in the studied agroforestry system present a great potential to prevent soil erosion and rebuilt soil C content. But, most of the residues, if decompose alone, would not supply sufficient nutrients, and need to be, therefore, mixed with leave of other species.

124
Table 5. Mean percentage of added C evolved as CO 2 of the tree- and bush-pruning residues from the shaded coffee system in southeastern Brazil, the mass remaining after 120 days of incubation at 25 C, and the results of a CENTURY model simulation using the same initial data Plant species % C-CO 2 Evolved Experimental Simulated MD s = 14.8 19.0 21.5 17.3 18.0 18.5 20.8 18.7 18.6 18.6
c

Mass remaining (% of original) Simulated MD s = 8.8 14.0 15.4 10.3 13.0 13.8 15.4 13.7 13.5 13.7
c

Experimental

Simulated MD s = 14.8 54.7 50.9 58.1 55.1 54.5 51.6 54.4 53.9 54.0

Simulated MD s = 8.8 74.3 71.1 80.3 75.5 74.6 71.1 74.2 74.3 74.1

C. cajan S. variable C. fernuginea P. gonoacantha C. urucurana M. multiora Mixture 1 Mixture 2 Mixture 3

13.7 14.2 10.9 12.6 13.3 15.0 13.1 12.6 13.0

0.9 a c b 0.9 b 0.5 f 0.7 e 0.8 c,d 1.0 a 0.8 d,e 0.7 e 0.8 d,e

74.8 72.7 82.2 76.8 75.0 72.0 74.6 77.3 74.8

3.0 2.3 4.6 4.4 3.3 1.8 2.9 4.1 2.8

b,c c,d a b b,c d b,c b b,c

a Numbers represent the mean value with standard deviation. bMeans within a column with different letters indicates a signicant different at the P< 0.05 level (n = 6). cMD s is the constant representing the maximum surface metabolic decomposition rate.

Acknowledgements We thank Paulo do Amaral Lopes, Donizete Lopes and Joo dos Santos for allowing us access to their farms for this study. We also thank Jos Braz Junior for expending extra time preparing samples and doing part of the pruning characterization. We would like to thank Dr. William Parton for discussions regarding CENTURY, and Ms. Robin H. Kelly for her technical assistance with CENTURY. This study was supported by a grant from CAPES, Brazilian PostGraduate Federal Agency. The mention of specic instruments or products is for the readers information only, and does not imply endorsement by the USDAARS or the Universidade Federal de Viosa.

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