Agroforestry Systems 61: 221236, 2004. 2004 Kluwer Academic Publishers. Printed in the Netherlands.

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Ecological interactions, management lessons and design tools in tropical agroforestry systems
L. Garc a-Barrios1, and C.K. Ong2
1 El

Colegio de la Frontera Sur, M exico. Carretera Panamericana y Perisur (s/n); San Crist obal de las Casas, Chiapas, Cp. 29290 M exico; 2 World Agroforestry Centre, P.O. Box 30677, Nairobi, Kenya; Author for correspondence: e-mail: lgarcia@sclc.ecosur.mx

Key words: Growth resources, Indices, Predictive understanding, Roots, Simulation models, Treecrop interactions

Abstract During the 1980s, land- and labor-intensive simultaneous agroforestry systems (SAFS) were promoted in the tropics, based on the optimism on tree-crop niche differentiation and its potential for designing tree-crop mixtures using high tree-densities. In the 1990s it became clearer that although trees would yield crucial products and facilitate simultaneous growing of crops, they would also exert strong competitive effects on crops. In the meanwhile, a number of instruments for measuring the use of growth resources, exploratory and predictive models, and production assessment tools were developed to aid in understanding the opportunities and biophysical limits of SAFS. Following a review of the basic concepts of interspecic competition and facilitation between plants in general, this chapter synthesizes positive and negative effects of trees on crops, and discusses how these effects interact under different environmental resource conditions and how this imposes tradeoffs, biophysical limitations and management requirements in SAFS. The scope and limits of some of the research methods and tools, such as analytical and simulation models, that are available for assessing and predicting to a certain extent the productive outcome of SAFS are also discussed. The review brings out clearly the need for looking beyond yield performance in order to secure long-term management of farms and landscapes, by considering the environmental impacts and functions of SAFS. Introduction Traditional low-input agricultural systems involving trees have been designed and managed for centuries by poor peasants around the world, and are still conspicuous in the tropics. During the past century, landuse intensication, agroecosystem simplication and other social changes have undermined the functionality of many of these low-input systems, and confronted peasant agriculture with enormous sustainability challenges (Nair 1998; Garca-Barrios and Garca-Barrios 1992; Garca-Barrios 2003). In the past two decades, great expectations have been set on the promotion of traditional and novel agroforestry practices as a means for slowing down or reversing such trends, once it became clear that high-input strategies promoted by development agencies had failed to be adopted and/or to deliver benets to smallholders (Sanchez 1995). Where there is still scope for fallow agriculture in the tropics, sequential agroforestry systems such as enriched fallows have been proposed; where land-use intensication and fragmentation is more severe, the bet has been on simultaneous agroforestry systems (SAFS) such as alleycropping, alley farming, parkland systems and trees on eld boundaries. During the 1980s, alleycropping was promoted throughout the tropics as a sustainable option for lowinput agriculture. By the 1990s it was recognized that the density, management intensity and environmental scope of new tree-based SAFS had been pushed too far. It became clear that introducing trees in croplands was in some cases like walking on a razor-edge because trees provide peasants with both crucial products and strongly facilitate crops, but can

222 also exert very strong competitive effects. Among other things, this motivated researchers to unravel the science of agroforestry (Sanchez 1995). The aim was a predictive understanding (at the ecological, physiological and agronomic level) on how such interactions and their productive outcome respond to species selection, tree management practices and resource limited environments, in order to develop realistic agroforestry solutions. In the process, a number of useful concepts, predictive models and production assessment tools were enhanced or developed. At the same time it became clearer that socioeconomic motivations and restrictions strongly inuence how farmers perceive and manage treecrop interactions, and that recommendations based on productivity at the plot level alone are insufcient for the long-term management and sustainability of farms and landscapes. This chapter addresses the issues previously mentioned and reviews current advances in treecrop interactions and their implications, focusing on annual crops and trees in simultaneous agroforestry systems in the tropics. Two companion chapters in this volume, by Jose et al. (2004) and Thevathasan and Gordon (2004), discuss interspecic interactions in temperate agroforestry systems. Until recently, ecologists accepted niche differentiation theory (MacArthur and Levins 1967) as the most important explanation of plant coexistence, and agroecologists consequently predicted that there must be considerable scope for obtaining or enhancing mixed crop over-yielding by nding the proper species combinations, spatio-temporal designs and simple management tasks which produce minimum interspecic competition (e.g., Vandermeer 1989). Ecologists have realized that weak interspecic competition is only one of many conditions, which can explain species coexistence in a natural plant community (GarcaBarrios 2003). Agroecologists and agroforesters have also found that complementarity is less conspicuous and requires more plant management than originally thought (Ong and Leakey 1999). A plant facilitates another plant when it modies the biophysical environment in such a way that it creates one or more potential conditions for a better performance of the latter (Hunter and Aarssen 1988). The facilitator can a) produce a net resource increase in the system, b) enhance the potential capture and/or use efciency of one or more growth resources, c) produce hormone-type stimulatory or inhibitory allelopathic compounds and/or d) deter the presence of competitors, predators and pathogens and e) attract pollinators or dispersal agents (Holmgren et al. 1997). It is important to stress that a) such potential benets can have little actual facilitative effect when other nonfacilitated resources or conditions limit growth (Kho 2000), and b) that the facilitators competitive and other negative effects can override its positive effects. The interplay of positive and negative plant interactions and its outcome can change along resource gradients, both in natural communities (e.g. Pugnaire and Luque 2001), intercrops (e.g., Rao and Willey 1980) and SAFS (e.g., Rao et al. 1998). Predicting the consequence of a change in environment is elusive, as results are highly variable and trends contradictory. A clearer and more comprehensive picture is beginning to emerge as researchers consider to what degree each resource is limiting to growth in an environment and how the species involved are suited to compete for and/or to facilitate the use of such resource(s) (Kho 2000; Ong et al. 2004). Treecrop interactions in SAFS share the basic features indicated in the previous section. Yet, trees have noteworthy characteristics that make them have very strong positive and negative effects on agroecosystem structure and function (Ong 1995). Trees have more biomass, are taller, and have more extended and

Positive and negative tree effects on crops in SAFS Competition occurs when two overlapping plants reduce one or more growth resources to the point where the growth, survival or reproductive performance of at least one of them is negatively affected (Harper 1990). Overlap increases as a consequence of growth and/or increased density. Plants differ greatly in size, life form, phenology and capacity to capture and use efciently above- and below-ground resources (Goldberg 1990); therefore, intra- and interspecic competition can differ strongly. When acting together in plant mixtures, these two interactions can produce a range of situations. At one extreme, species coexist and overyield (i.e. more land would be required to obtain the mixture yields by sowing each species separately as sole crops). At the other, a single dominant species strongly reduces the performance of others. When above- and/or belowground resources are partitioned between trees and crops such that relative interspecic competition is lower than relative intraspecic competition, we refer to the case as weak interspecic competition, complementarity or niche differentiation (Vandermeer 1989).

223 deeper roots than crops. Trees are perennial and therefore: a) they eventually explore a relatively large space and can substantially modify their biophysical environment to their benet, b) they are better adapted to resist cyclic environmental harshness and to use and recycle resources when it is more efcient to do so, and c) they commonly have a canopy and a root system in place when crops starts growing (Ong et al. 1996). Mature trees in SAFS can have the following positive effects on crop elds: 1. They can add considerable amounts of organic matter to the soil and slow down its decomposition rate, improving soil fertility and physical structure. Trees roots can stabilize loose soil surfaces, which together with tree litter cover, reduces erosion (Young 1997). They recover leached nutrients from deep soil or bedrock layers inaccessible to crops (Rao et al. 1998). Trees can make more phosphorus available via mycorrizha and x important amounts of nitrogen, which can be transferred to the crops via shoot and root prunings (Giller 2001). 2. Although trees can increase the potential soil waterholding capacity, they have variable and conicting effects on the actual water volume available in the treecropsoil system: Rainfall intercepted by the canopy that evaporates without reaching the soil can be as much as 50% when tree density is high (Ong et al. 1996). Yet, reduced soil evaporation by tree shade can help offset such losses as long as rainfall is lower than 700 mm per annum (Ong and Swallow 2004). Trees can eventually increase inltration but this strongly depends on slope and soil characteristics (Ong and Swallow 2004). In general, tree presence produces a net increase in total water used by the system (Ong and Swallow 2004). 3. Tree shade can reduce leaf temperature and evaporative demand experienced by crops, increasing the latters water productivity (i.e., g of dry mass per g of water transpired). The net shade effect is more positive (or less negative) when the annual crop is a C3 plant which is normally light saturated in the open, so partial shade may have little effect on assimilation or even be benecial (Ong 1996). This improvement in conversion efciency is relatively modest when compared to the effect described in the previous point (Ong and Swallow 2004). 4. Tree hedgerows provide protection against wind and runoff (Rao et al. 1998.). 5. Trees can reduce weed populations and change weed oristic composition towards less aggressive, slow growing species (Leibman and Gallandt 1997). 6. Little is known about the complex and sometimes conicting effects of trees on pest and disease control, but Schroth et al. (2000) and Rao et al. (2000) provide comprehensive reviews on the topic. Increased pest and disease incidence has often been observed directly at the treecrop interface. Trees increase air humidity, which favors microorganisms, provide shelter for herbivores (insects, birds, and small mammals), which damage the crops, and reduce pest and disease tolerance of competition-stressed crops. Trees themselves can be more susceptible to pest and disease attack when sown at densities and spatial arrangements uncommon to their natural environments. Yet, tree hedges have the potential to slow down windborne pests and diseases, to act as repellants, and to attract natural enemies; recent evidence is provided by Girma et al. (2000). On the other hand, with increase in density of trees, their size, and/or ability to capture resources in SAFS, they can exert strong competition for light, water and nutrients, and reduce annual crop yields beyond the interests of farmers if improperly selected and managed (Garca-Barrios 2003). Nevertheless, weak competition is possible under certain circumstances such as the following: 1. Tree roots can potentially reach below the crop root zone, and thus they can use water accumulated deeper in the ground when the crop is growing; after the crop is harvested, they can use whatever residual available water is found in the crop root zone; and they can use any additional rain which falls outside the crop growing season (Ong et al. 1996). It is important to stress that trees used in SAFS do not always have deep pivotal roots, and that mixed and supercial tree root architectures are common (van Noordwijk et al. 1996). Moreover, if water recharge below the root zone is infrequent and/or nutrients are supercial, most trees will tend to develop or redirect their roots to the upper soil layers (Rao et al. 2004) and only a few species will develop roots that can reach relatively deep water tables. Consequently, there seems to be less scope for vertical root complementarity than originally thought (Sanchez 1995; Ong and Swallow 2004). 2. Some deciduous tree species used in SAFS in semiarid regions such as Faidherbia albida exhibit reverse phenology: they produce their leaves and demand water only during the dry season, while their litter provides nutrients and their trunk and bare branches cast a light shade over crops during the rainy season (Rao et al. 1998).

224 3. Hundreds of different nitrogen xing leguminous trees are used in SAFS (Giller 2001). When their leaves are not used as mulch, they do not facilitate crops, but their alternative N source can signicantly reduce competition for this resource. 4. Radiation can be shared between closed canopy tree stands with low tree-LAI (leaf area index) and understory crops which tolerate or even require light shade, yet, most C4 and C3 annual crops are fast growing species which do not easily adapt to this condition (Black and Ong 2000). In most SAFS there are problematic tradeoffs between positive and negative tree effects. Trees can modify the soil and microclimate environment much more than crops; they can have strong facilitative effects; and they can produce important environmental services and in specic circumstances compete weakly for some resources. Yet, tree characteristics often confer them a clear competitive advantage and they can strongly out-compete crops or reduce crop yield beyond acceptable levels to the farmer. Design and management practices such as optimum density and spatio-temporal arrangement, fertilization, weeding, and proper shoot and root pruning can help reduce competition and/or enhance facilitation. Yet, developing novel and successful SAFS designs and management practices has proved more difcult than expected. This has led to a number of partial failures as well as to useful lessons (Ong 1994; Sanchez 1995; Rao et al 1998). Management difculties arise in SAFS for a number of reasons, including the following: 1. Many SAFS practices are labor intensive and require relatively high crop-yield-advantage over sole crops to be attractive to farmers. This economic limitation is aggravated when novel SAFS are designed and tree species for them are selected mainly for mulch production and soil protection, with little attention to more immediate and direct tree products such as fodder, lumber and wood (Ong and Leakey 1999). 2. Practices that reduce competition for a given resource can inadvertently increase competition for other resources and/or decrease facilitation. For example, in tropical alleycropping systems: a) Intensive shoot pruning and accumulation of N-rich mulch in the alley reduce light competition and facilitate N allocation to the crop; yet, they both promote lateralsupercial tree roots which strongly compete with the crop for water and nutrients, sometimes overriding the positive effects (Hairiah et al. 1992; Fernndez et al. 1993). b) Tree distances can be increased and optimized (Vandermeer 1998), and roots pruned (Ong and Swallow 2004), but in some cases with a negative effect on the formation of the root safety net (van Noordwijk et al. 1996). 3. The consequences of expanding successful SAFS to other environments have proved difcult to predict until recently, and the environmental scope of promising SAFS has been more limited than expected. The way in which tree-crop interactions respond to changes in environmental resource balance partially explains these situations. For example, alleycropping was extended to arid and semiarid regions, generally with very poor results (Rao et al. 1992). It was later realized that tree competition for water strongly depressed crop yields and was not compensated by the amounts and quality of mulch (Rao et al. 1992). Similar disappointments were obtained in very acid and unfertile soils (e.g., Ultisols and Oxisols), where P and other nutrients rather than nitrogen were the most limiting resources (Sanchez 1995). In order to optimize tradeoffs between positive and negative tree effects, it is necessary to select in each environment the proper annual crops, tree species and densities, and soil amendments. It is also important to correctly choose the timing and intensity of rootand shoot pruning. Proper design and management is critical to achieve the desired balance between tree growth, annual crop production, tree crop production, weed suppression, soil improvement and other environmental services. Among other things, this implies nding tree/crop combinations that promote resource facilitation, resource sharing, and minimum resource competition and are compatible with farmers goals and conditions. These combinations are not always readily obvious or easily found. A number of analytical and simulation tools have been developed in the past decade which help to understand some of these interactions and their productive outcome. We will now discuss some examples that represent the different approaches and their scopes.

Treecrop interactions and yield assessment Methods that do not consider growth resources explicitly Quantifying the net effect of interactions The combined effects of intra and interspecic interactions on species and community yields in mixtures can be quantitatively assessed without separating pos-

225 itive and negative effects explicitly, and without considering resource distribution and use. An extensive literature has been developed on this matter during the past 40 years (Vandermeer 1989). A number of competition- and productivity indices have been developed and their scope, limitations, and pitfalls discussed (Connolly et al. 2001; Williams and McCarthy 2001). Most productivity indices have been developed for two species intercrops. They assume that the farmer is interested in nding whether combinations of both crops in the same eld can outperform the corresponding sole crops. We will give a very brief account of how crop and tree yields relate separately to their net intra and interspecic interactions by developing a graphical analysis of a hypothetical and generic SAFS. We will then discuss the simultaneous assessment of crop and tree yields. Consider the crop and tree sole stands in Figure 1(a) and (b). Both are sown at their optimum sole crop densities (N.), i.e., the number of individuals per unit area which produces the maximum possible yield (Y.) in that surface. These parameters result from the reasonably hyperbolic relation between sole crop yields and sole crop plant densities (Willey and Heath 1969) depicted in Figure 1d and are a direct consequence of net intra-specic interactions. In our example, the sole crop stand parameters are Nc = 6 and Yc = 6, while the sole tree stand parameters are Nt = 2 and Yt = 18. Intra-row plant distances are the same for both crops. In the treecrop mixture depicted in Figure 1c, every sixth crop row has been substituted with a tree row. As a consequence, the crop density in mixture (nc) is 5 and the tree density in mixture (nt) is 1. In other words, nc = 5/6*Nc and nt = 1/2*Nt. For simplicity, we will assume that both tree competition and facilitation on the crop can be present, and that no tree pruning is practiced so that facilitation results from other mechanisms. Crop plants nearer to the tree are generally smaller as they experience more competition (see Figure 1c), but here we consider the average plant performance within the unit area. Different crop yield outcomes in mixture (yc) are possible; they are shown in Figure 2a. An interesting starting point is case 3, where yc = 5/6*Yc, i.e., mixture crop yield is reduced in the same proportion as crop density. The average crop individual has the same weight in sole crop and intercrop, which means that the average net effect of a tree on crop plants matches the net intra-specic effect of the average crop individual in the sole stand. Considering the asymmetry in tree and crop sizes, this seems highly unlikely, unless it results from very weak tree competition due to high resource complementarity, or from facilitation almost compensating competition. In case 4, either tree competition is zero (perfect complementarity) or facilitation exactly compensates competition, because in this case, yc equals the yield expected for ve plants per unit area in the sole crop (here we make the assumption that given a near-optimum crop density, particular plant arrangement has little effect). In case 5, the crop stand experiences enough net facilitation to match the maximum sole crop yield, and in case 6 to surpass it. In case 1 and 2 the average crop plant experiences signicant net competition and crop yields are strongly reduced. A similar analysis is possible from Figure 2(b), for the trees performance in mixture; the roles are simply inverted. Cases 1 and 2 are highly unlikely, unless strong crop allelopathy effects on young trees are present. The most probable outcome is somewhere between case 3 and 4. Cases 5 and 6 are not realistic, given asymmetry between tree and crop. Of course, the crop and tree yield outcomes in our example will be coupled and depend on one another such that not all outcome combinations make sense. Some possible combinations are depicted graphically in Figure 3. In more general terms, each specic spatio-temporal mixture design for a given pair of plant species in a given environment produces a pair of coupled yields. The set of coupled yield outcomes of all possible designs (which include all sole crop yields as well) is called a yield set (Vandermeer 1989); the subset of points which constitute the exterior envelope of this set is called the production possibility frontier (PPF; Ranganathan and De Wit 1996); (Figure 4 presents a possible yield set and PPF for our hypothetical SAFS). A PPF can be analyzed to compare sole and mixed crop outcomes and to nd the optimum mixed crop designs according to different biological and economical performance criteria. An example of such criteria is the land equivalent ratio (LER), dened as (yc/Yc) + (yt/Yt). A LER > 1 means that there mixture is advantageous because more land would be required to obtain yc and yt by sowing each species separately as sole crops. (For further details on LER and other mixture performance indices, see Vandermeer 1989). A few scores of yield pairs can be obtained through eld experiments, while more thorough yield set and PPF constructions can be aided by experimentally tted models. Simple analytical models have been developed for this purpose, which only consider global population densities and render nal

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Figure 1. Schematic representation of the yield vs. density relation of a tree-crop mixture, and its corresponding sole crops stands. Bar areas represent the aboveground biomass of the individual plants. (a) sole crop stand; (b) sole tree stand; (c) tree-crop mixture; (d) sole stand yield vs. density relations.

Figure 2. Sole stand yield vs. density relations and some possible yield outcomes (16) after mixture with the other species. (a) Crop yields; (b) Tree yields. See text for further explanation of mixture outcomes.

227 biomass or yield (e.g., Ranganathan and De Wit 1996). Spatially explicit, individual-based plant growth models have also been developed. They require empirical growth and competition parameters and render yields at any growth stage for any spatio-temporal design (e.g. Garca-Barrios et al. 2001). Both models need to be re-parameterized in different environments. Quantifying the effects of positive and negative interactions separately The idea to separate and quantify positive and negative tree effects on crop yield in SAFS was formalized by Ong (1995) in the simple equation: I=F+C (1) high; 3) positive and negative component effects are very site specic and change with the environment . After a modication by Ong (1996), the equation evolved to (Rao et al. 1998): I=F+C+M+P+L+A (3)

where I is the overall interaction, i.e., the percentage net increase in production of one component attributable to the presence of the other component, F is the fertility effect, i.e., the percentage production increase attributable to favorable effects of the other component on soil fertility and microclimate, and C is the competition effect, i.e., the percentage production decrease attributable to competition with the other component for light, water and nutrients. The I value is based on total area, including that occupied by trees; therefore, a positive I value means net increase in total crop yield, irrespective of crop population in mixture relative to that of sole crop (Rao et al. 1998). In alleycropping systems, the measurement of F and C has been accomplished with four treatments: Co = sole crop; Cm = sole crop + mulch from pruned trees; Ho = crop + tree with mulch removed; Hm = crop + tree with its mulch. The equation then becomes I = (Cm Co) (Hm Cm) (2)

where F refers to effects on chemical, physical and biological soil fertility, C to competition for light, water and nutrients, M to effects on microclimate, P to effects on pests, diseases and weeds, L to soil conservation and A to allelopathy effects. The benet of Equation (3) is that it encapsulates a comprehensive overview of the possible effects involved. However, as emphasized by the authors, many of these effects are interdependent and cannot be experimentally estimated independently of one another. Limitations have been identied for this approach (Ong et al. 2004): 1) due to interdependence, the individual terms most likely will give a sum that exceeds I, such that the relative importance of each term cannot be established. 2) It cannot predict delayed effects and long-term trends. 3) It is not meant to predict the consequences of moving from one environment to another, as it does not explicitly consider growth resource capture and use, and their interaction (e.g. water x P interaction in Pxing soils). Cannell et al. (1996) attempted to clarify the resource base of Ongs equation but did not make plant interaction with resources sufciently explicit. Mechanistic research may be necessary to understand SAFS functioning and performance over time and/or in different environments. Methods that explicitly consider growth resources Positive and negative interactions between plant species largely depend on how the latter affect each others ability to capture and use growth resources. The principles of light, water and nutrient capture and use efciency rst applied to sole crop growth analysis and modeling have been extended to intercropping and agroforestry research in the past decade. The eld has seen enormous advances in the gathering of relevant experimental data, theory development, modeling capability, and construction of very sensitive instruments for measuring direct above- and belowground resource ow and capture (Black and Ong 2000). They have also made evident the challenges for studying these processes in multispecies systems, and the limitations of some basic idealized and simplied assumptions about the relation between plant growth and resource capture and use efciency.

The competition term can also be calculated as (Ho Co) or, more conveniently, as the average of (Hm Cm) and (Ho Co). This equation motivated the analysis of numerous previous alleycropping experiments and the establishment of new ones (Ong 1996). Unfortunately, few proved to have these four treatments and/or the appropriate experimental designs and eld display. Nevertheless, successful positive and negative separation in a few experiments made evident, among others, the following important facts (Sanchez 1995; Ong 1996; Rao et al. 1998; Ong et al. 2004): 1) Strong tree competition is more conspicuous than originally thought; 2) high tree growth rates and tree biomass, intuitively associated with the potential for a signicant fertility effect are also strongly related with tree competitiveness, so tradeoffs between both interactions are

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Figure 3. Coupled tree and crop yields in mixture. Points 16 represent the six crop yield cases depicted in gure 2(a), and reasonable associated tree yields. Three additional outcomes are included: in A, tree excludes crop; In B crop excludes young tree through strong allelopathy. In C allelopathy effect on tree is milder. The slope of the dashed line indicates the intensity of the slightly negative crop effect on tree biomass. Stars are maximum sole crop yields, and the line that crosses them represents all outcomes with a land equivalent ratio (LER) = 1. Above this line, LER > 1.

A central tenet of mechanistic plant-plant interaction analysis has been that, according to the Law of the Minimum (Blackman 1905), as long as a resource is the most limited, growth depends linearly on the capture of this resource. If eventually another resource becomes the most limiting, the formers concentration ceases to have an effect and the latter dictates growth. As a consequence, biomass production (W) should be easily modeled as the product of the capture of the most limiting resource, and the efciency with which the captured resource is converted into biomass (Monteith et al. 1994; Ong et al. 1996). The conversion efciency of the limiting resource is considered to be conservative for a given species for a given environment, and the growth response of the plant is attributed to the increased capture of the resource. Research data (e.g., Demetriades-Shah et al. 1992; Black and Ong 2000) and theoretical developments (Kho 2000; Ong et al. 2004) have shown that: 1) Usually several resources are limiting, in which case the relation between biomass production and the capture of one single resource should be viewed as a correlation, not a causal relation that can be readily modeled. 2) The

variation in conversion efciencies between environments is larger than that between species; therefore such efciencies are more determined by the environment than by species. 3) Plants alter the availability of resources simultaneously and thus conversion efciencies by changing resource limitation. Moreover, the most limiting resource for a species can differ in sole system and mixture. 4) Efciencies should be studied and modeled in relation to the availabilities of the other resources, and treated as variables in process-based models. 5) Dynamic simulation models for different resources should be linked. In recent years, different resource-based modeling approaches have been developed and/or used to explore or predict how tree-environment-crop interactions (and the productive performance of SAFS) change when environmental resource availability is modied. Some relevant examples are cited below: The mulchshade model Tree canopies in alleycropping provide N-rich mulch but reduce radiation available for crops. The net tree effect should then be a function of at least three factors: 1) the trees mulch:shade ratio (MSR), 2) the

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Figure 4. A yield set and Production Possibilities Frontier for the hypothetical tree-crop mixture presented in gures 1 (c). Thin solid lines result from xing a global density and modifying the crop-tree proportion; thick dashed line from xing a tree-crop proportion and modifying global density. The thick solid line is the production possibility frontier, and the area under it is the yield set. The thin dashed line represents all outcomes for which LER = 1.0, and the black dot on the PPF is the tree crop mixture which renders the highest attainable LER value.

importance in the particular environment of nitrogen supplied by mulch, expressed as a Nsoil : Nmulch ratio (NNR), and 3) tree row distance as a surrogate of tree leaf area index. Van Noordwijk (1996) developed a static-analytical alleycropping model, which links these species attributes and management factors, and produces explicit algebraic solutions. The simple MSR offers a basis for comparing and selecting tree species. The equation variables and parameters require a lengthy explanation; here we are interested only in discussing the type of results it delivers. The model predicts that at low soil fertility, where the soil fertility improvement due to mulch can be pronounced, there is more chance that an agroforestry system improves crop yields than at higher fertility where the negative effects of shading will dominate. Moreover, it denes combinations of MSR and NNR for which some alleycropping systems should be expected to work (Figure 5a). Although not validated with empirical data, it has been parameterized for typical alleycropping tree species in order to estimate their optimum hedgerow density at different Nsoil values

(Figure 5b). The mulch/shade model provides useful insights, but does not incorporate the interactions between resource dynamics, and crop and tree growth. Incorporating these elements extends the model beyond what can be solved analytically and into the realm of dynamic simulation models. Integrated water, nutrient and light simulation models in agroforestry systems In the late 1990s complex agroforestry simulation models and user platforms were constructed for example, WaNuLCAS (van Noordwijk and Lusiana 1999) and HYPAR (Mobbs et al. 2001), and they are still undergoing development, parameterization, and validation. These two models are process-based and have a useful level of spatial structure. Both require weather databases and a considerable number of soiland plant parameters as inputs. WaNuLCAS is quite elaborate in its soil sub-models and HYPAR in its canopy sub-models. They are too complex to allow a useful description here, but excellent specications and user manuals are available. Once proper inputs and

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Figure 5. Some of the results obtained by van Noordwijk, (1996) with the mulch and shade model. (a) shows combinations of MSR and NNR for which some hedgerow systems should be expected to work, given two different values of N mulch. P, C and L stand for estimated MSR values for Peltophorum dasyrrachis, Calliandra calothyrsus and Leucaena leucocephala, respectively. (b) shows predicted optimum tree density in an alley cropping system for ve tree species (Peltophorum dasyrrachis, Erytrina poeppigiana, Gliricidia sepium, Leucaena leucocephala and Calliandra calothyrsus) as a function of Nsoil. Tree density is expressed in linear meters of hedgerow per hectare. Species parameters were estimated empirically and are reported in van Noordwijk, (1996), table 3.2.

parameters are in place, these models can be powerful tools for systematically exploring the possible consequences of diverse management practices and of one or more resource gradients, before engaging in costly and time consuming experiments and productive projects. Van Noordwijk and Lusiana (1999) used WaNuLCAS to model grain- and wood production, water use, water-use efciency and N limitation for a wide range of annual average rainfall conditions (240 mm to 2400 mm) in parkland systems with differently shaped trees; an example of their results is presented in Figure 6a and 6b. As yet, no experimental data set exists on the same agroforestry system at the same soil but widely differing rainfall conditions, so they used data and parameters from different sources and from theoretical assumptions. Model results generally agreed with conclusions derived from experimental evidence (Breman and Kessler 1997). Similar conclusions were reached with the HYPAR model about the effect of climatic gradients on treecrop interactions and productive outcomes (Cannell et al. 1998). Complex process-based models are potentially powerful tools but have their own limitations and pitfalls; they are expected to confront problems of parameter estimation, validation and input gathering

similar to or more severely than those found in sole crop physiological models (Aggarwal 1995; Hakanson 1995). These models can be used with caution in order to gain insight about management practices and about the consequences of theoretical assumptions; their outputs should be constantly confronted with empirical results and farmers experience. A general treeenvironmentcrop interaction equation for predictive understanding of SAFS Kho (2000) has developed a simple but powerful analytical model in which the overall tree effect on crop production is explained as a balance of (positive and negative) relative net tree effects on resource availability to the crop. We will describe here some of its basic features, consequences and applications. When trees are introduced in a crop eld, they simultaneously change the availability of several resources in the environment of the crop, some for the better and some for the worse. Notwithstanding the Law of the Minimum stated earlier, it has become clear that not one but many resources can limit growth simultaneously and that the degree to which a resource affects production at a given level is dependant on the availability of the other growth resources (Kho 2000). Consequently, the relation between a given resource and production

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Figure 6. Some of the results obtained by van Noordwijk and Lusiana (1999) with the WaNuLCAS model for grain and wood production, for a range of annual rainfall conditions in an hypothetical agroforestry system where trees provide N-rich mulch. For comparison, the sole crop and sole tree stand are grown also. (a) shows a gradual shift from water to N as the major factor limiting crop production as rainfall increases. At low rainfall levels, tree competition for water dominates over positive effects of N supplied by tree mulch. (b) Because of crop competition, tree yield is lower in the SAFS stand than in sole tree stand.

is not linear but asymptotical when other resources are held constant. As a particular resource becomes less available in relation to others, its inuence on production becomes greater and in this last sense it becomes more limiting. The limitation of a resource Ai (i.e., Li) can be formally dened as the ratio between the slope of the production response curve (at a given resource level) and the average use efciency of that level of resource by the crop; it is therefore a relative non-dimensional term. Li can also be dened (more intuitively) as the relative change in production in response to a relative change in resource availability (Kho 2000), which corresponds to the denition of elasticity in economic theory. Kho 2000 has fruitfully applied Eulers law from economics to demonstrate that the sum of limitations (or elasticities) of all growth resources ( Li) should reasonably be equal to 1.0 if the so called constant return to scale assumption holds. De Wit (1992) shows agricultural data supporting this latter assumption of proportional relation of output to input.

As a consequence of the previous arguments, when trees interact with a crop, the expected relative crop yield change (dW/W) should be equal to a weighted sum of the relative changes induced by the tree on each resource Ai; the weights should be the Lis which result from the particular balance of resources in the specic environment. This leads to the equation:
n

dW/W =
i =1

(dAi/Ai) Lii

(4)

In short, each resource contributes to the relative change in production proportionally to its degree of limitation and proportionally to its relative change in availability. From Equation (4), Kho (2000) derives Equation (5), which predicts the relative change in crop production (I, as in Ong 1995), as a function of tree effects on resource availability and of resource balance in the particular environment considered:
n

I =
i =1

Li Ti

(5)

232

Figure 7. Trees inuence crop production through altering (the balance of) resource availabilities to the crop. Each rectangle represents a different resource: water (W), nitrogen (N), phosphorus (P) and radiation (R). The height of each shaded area relative to the height of the rectangle represents the relative change in availability of the resource (Ti). The width of each shaded area relative to the total width represents the limitation of the resource in the tree-crop interface (Li). The sum of positive and negative shaded surfaces relative to the total surface of the rectangle represents the overall tree effect I expressed as fraction of sole crop production. The gures show possible tree effect balances of an alley cropping technology in a humid climate. A) on nitrogen decient soils, B) on acid (phosphorus decient) soils, C) on nitrogen decient soils with nitrogen fertilizer (applied to the alley crop and the sole crop), and D) on acid soils with phosphorus fertilizer (applied to the alley crop and the sole crop). The relative net tree effects on availability of each resource (Ti) are equal in AD; only the environments (i.e. resource limitations Li) change, and explain the different overall effects (I). Source: Ong, et al., 2004.

where Ai Ai;multi Ai;mono = (6) Ti = Ai Ai;mono Ti is the relative net change in availability of resource i because of the tree, Ai;multi is the availability of resource i to the crop in the SAFS and Ai;mono in the sole crop. Robust estimations of L and T values for a particular SAFS and environment can be obtained experimentally and/or derived from the literature following relatively simple methods which are described and applied by Kho (2000) and Kho et al. (2001). The relation between resource balance and limitation combined with Equation (5) leads to two rules

that can be viewed as counterparts of classic crop production principles (Kho 2000): 1) The greater the availability of a resource in the environment, the smaller is its share in the overall tree-crop interaction. 2) The greater the availability of other limiting resources in the environment, the greater is the share of a resource in the overall tree-crop interaction. These rules are helpful for predicting the performance of a SAFS technology when it is extended to another environment and for developing a SAFS technology. For example, Kho (2000) showed that for the alleycropping technology the net effect of the alleys on the availability (to the crop) of the resources light, water, and phos-

233 phorus is most likely negative, and that for nitrogen it is most likely positive. Consequently, in a (sub-) humid climate on nitrogen- decient soils, the overall alleycropping effect is most likely positive, because of the high limitation for the positive nitrogen effect and the low limitations for the negative net effects on other resources (Figure 7A). In the same climate, but on acid soils, phosphorus is relatively less available, which will increase the share of the negative phosphorus effect (rule 1) and will decrease the share of the positive nitrogen effect (rule 2), resulting in a negative overall effect (Figure 7b). Through management practices, the share of positive net effects can be increased and the share of negative net effects decreased. Compare, for example, Figure 7b and 7d for the effect of phosphorus fertilizer applied to both the alleycropping system and the sole crop system. External inputs of organic or inorganic nitrogen are most likely inappropriate, because these will reduce the share of the positive nitrogen effect (rule 1) and increase the share of the negative effects (rule 2; cf. Figure 7a and 7c). production and resource use. The principal agroforestry systems suited to humid tropical environments are multistrata systems, perennial crop combinations, managed tree fallows, contour hedgerows and reclamation agroforestry (ICRAF 1996; Young 1997; Tomich et al. 1998). These mixtures of trees and crops have the potential to improve land management via their ability to reduce soil erosion and improve soil conditions for plant growth. There is some evidence for the utility of agroforestry systems for soil conservation, soil organic matter maintenance, nutrient retrieval, and nutrient recycling (Buresh and Tian 1998; Young 1997). In farming for annual crops, contour hedgerows may provide a viable alternative to conventional conservation measures (Kiepe and Rao 1994). However, despite the demonstration that such systems can dramatically reduce soil losses and improve soil physical properties, the benecial effects on crop yield are often unpredictable and insufcient to attract widespread adoption (Alegre and Rao 1996). Agroforestry is now receiving increasing attention by researchers, landowners and policy makers in Australia as a potential solution to the salinity problems caused by the rising water-table (Lefroy and Stirzaker 1999). Replacement of the native vegetation, mainly trees and shrubs, have resulted in a steady increase in the water table for much of the semiarid wheat (Triticum sp.) belt of Eastern and Western Australia, because the vegetation has been replaced by wintergrowing crops such as wheat (Triticum aestivum), barley (Hordeum vulgare), canola (Brassica napus), and lupin (Lupinus spp.), which cannot fully utilize the annual rainfall. The salinity problem is more complex than that experienced in the tropics because of the nature of the duplex soils there, which slow lateral movement of water across the landscape and there are few protable tree species to replace the existing annual crops. Widely spaced tree rows (10 mm to 300 m and also called alleycropping) of fast-growing native (e.g., Eucalyptus spp.) and exotic origin (e.g., tree lucerne Chamaecytisus palmensis) were originally seen as the practical alternative for solving this huge landscape problem. As with the tropical alleycropping experience, there is a strong tradeoff between environmental function and crop performance in the Australian environment. Therefore, there is now a serious emphasis on identifying trees that can provide direct value to farmers (e.g., oil malle, Eucalyptus polybractea), as suggested by Ong and Leakey (1999) for sub-Saharan Africa, in addition to the hydrological function.

Beyond yield performance During the past two decades, agroforestry research in the tropics has focused, naturally, on the prospects for higher productivity in low-input agriculture, which are based on plot-level research just as in agronomic research. In recent years, there is a growing realization that recommendations based on productivity at the plot level alone are insufcient for the long-term management of farms and landscapes. Sustainability of land use practices, a major incentive for preferring agroforestry to high input agriculture, depends (among other things) on ensuring that the ow of energy through the agroecosystems is in close balance to those of the natural ecosystems (Lefroy and Stirzaker 1999). Furthermore, extrapolation of results from plot to landscape level is often awed because they fail to account for the lateral movement of water and soil, which are greatly inuenced by lters in the landscape (van Noordwijk and Ong 1999). Young (1998) argues that much of the future increase in food and wood production in the humid tropics will have to be achieved from existing land and water resources. Therefore, future research agenda should aim to improve the efciency with which land and water are currently used. One promising option for improvements of this kind is by using agroforestry, with the ultimate aim of achieving sustainability of

234 Conclusions and future research needs During the past decade it was recognized that introducing trees in croplands to promote sustainable low-input SAFS in the tropics was a challenging task. Some reasons for this are: a) trees provide both crucial products for smallholders and strongly facilitate crops, but can also exert stronger competitive effects than previously expected; b) practices aimed at increasing trees benecial effects can sometimes also enhance trees competitiveness; c) the interplay between positive and negative effects of trees change sometimes signicantly from one environment to another. This makes it difcult to predict the consequences of extending successful agroforestry practices to new environments having different resource levels and resource balances. This motivated scientists to develop predictive understanding of how such interactions and their production outcome respond to species selection, tree management and resource availability in order to develop realistic SAFS solutions. Among other research tools and strategies, a number of analytical and simulation tools were developed and/or adapted during the period in order to assess tree and/or crop yields. A rst group of static- or dynamic models analyze the yield outcomes of positive and negative tree effects without considering growth resources explicitly. A second group does consider these and can therefore predict to a certain extent the effect of the growth environment on tree-crop interactions. Static and dynamic models that do not explicitly consider resources are comparatively easy to parameterize and to use in determining tree-crop interaction effects (I) and construction of production possibility frontiers, and can shed light on specic questions, but need to be calibrated for each particular environment. Spatially explicit, processbased, simulation models (which balance resource use explicitly) are comprehensive and quite powerful research and design tools for SAFS, but are difcult and costly to parameterize. Non-modelers might make a more efcient and better-targeted use of them by acquiring insights provided by simple but powerful analytical models such as the tree-crop-environment equation described previously. Model outputs should be constantly confronted with empirical results and farmers experience, needs and constraints. It is also important to bear in mind that recommendations based on productivity at the plot level alone are insufcient for the long-term management and sustainability of farms and landscapes. The partial failure of labor-intensive agroforestry practices using very competitive fast-growing trees at relatively high densities is re-directing SAFS research. Where light-demanding annual crops are important to farmers and soil resources are limited and/or fragile, major efforts should be directed to low tree-density SAFS. Tree species selected for this purpose should provide both environmental services and immediate products and be amenable to pruning during low labordemand periods. In more general terms, a exible approach to SAFS design need to be developed with tree density and management as a function of a) environmental and economical constraints and b) relative value of crop and the tree products and services. In the tropics, where drylands and/or unfertile soils are predominant, competition for water and nutrients is critical. Belowground tree-crop interactions and the effects of root pruning strategies are receiving more attention and their research especially to develop economically usable root pruning practices should be pursued further. Tree effects on weeds, herbivores and pathogens, which reduce crop yields, need more attention and their effects should be incorporated into quantitative assessment analysis. SAFS (and all agroforestry systems) are attracting increasing attention as one of the potential mitigations for large-scale soil problems and negative effects of global warming. It is therefore necessary to consider the environmental impacts and functions of SAFS at a broader scale. Similarly, the long-term effects of SAFS on soil and water resources should receive attention particularly in different soil and environmental conditions. Modeling efforts should continue. Linking plant growth processes and modeling approaches, designing inexpensive resource-ow measuring instruments and developing experimental designs and robust parameterization techniques seem to be the major challenges. Relatively simple equations and concepts can be used to unravel positive and negative aspects of treecrop interactions and how they inuence overall production in simple agroforestry systems. However, designing more complex multispecies SAFS with appropriate management practices presents additional challenges both for satisfying immediate household needs and for replication at different scales while achieving sustainability in the face of environmental and social change. Managing such systems has to be based more on monitoring, diagnosis, remediation, mitigation and adaptation, rather than on a blueprint predictability of the behavior of the agroecosystem. The potential of process-based integrated resource use models in

235 designing and evaluation of complex systems needs to be tested. Research methods are needed to derive useful and exible rules from adaptive management.
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