Effects of species and soil‐nitrogen availability on root system architecture traits – study on a set of weed and crop species

Better managing crop : weed competition in cropping systems while reducing both nitrogen and herbicide inputs is a real challenge that requires a better understanding of crop and weed root architecture in relation to soil‐nitrogen availability. An original approach was used which considered the parameters of a simulation model of root architecture as traits to analyse (a) the interspecific diversity of root system architecture, and (b) its response to soil‐nitrogen availability. Two greenhouse experiments were conducted using three crop and nine weed species grown at two contrasted concentrations of soil‐nitrogen availability. Plant traits were measured to characterise both overall plant growth and root architecture, with a focus on primary root emergence, root elongation and branching. The studied root traits varied among species (from a twofold to a fourfold factor, depending on the trait), validating their use as indicators to analyse the interspecific variability of root architecture. The largest interspecies differences were for two traits: ‘maximal apical root diameter’ and ‘interbranch distance’ (distance between two successive laterals on the same root). Conversely, most of the studied root traits varied little with soil‐nitrogen availability (from no variation to a 1.1‐fold factor, depending on the trait) even though soil‐nitrogen availability varied with a 17‐fold factor and impacted the overall shoot and root biomass. So, the root traits used in this article are stable whatever soil‐nitrogen availability. As they reflect processes underlying root system architecture, this low effect of nitrogen suggests that the rules governing root architecture are little affected by plant nitrogen status and soil‐nitrogen availability. We propose that the determinants of differences in root system architecture between soils with contrasted nitrogen availability mainly originate from differences in the amount of carbon allocated to and within the root system. Characterising each plant species by a combination of root traits gave insights regarding the potential species competitive ability for soil resources in agroecosystems.     

A Mechanistic Study of Plant and Microbial Controls over R* for Nitrogen in an Annual Grassland

Differences in species'' abilities to capture resources can drive competitive hierarchies, successional dynamics, community diversity, and invasions. To investigate mechanisms of resource competition within a nitrogen (N) limited California grassland community, we established a manipulative experiment using an R* framework. R* theory holds that better competitors within a N limited community should better depress available N in monoculture plots and obtain higher abundance in mixture plots. We asked whether (1) plant uptake or (2) plant species influences on microbial dynamics were the primary drivers of available soil N levels in this system where N structures plant communities. To disentangle the relative roles of plant uptake and microbially-mediated processes in resource competition, we quantified soil N dynamics as well as N pools in plant and microbial biomass in monoculture plots of 11 native or exotic annual grassland plants over one growing season. We found a negative correlation between plant N content and soil dissolved inorganic nitrogen (DIN, our measure of R*), suggesting that plant uptake drives R*. In contrast, we found no relationship between microbial biomass N or potential net N mineralization and DIN. We conclude that while plant-microbial interactions may have altered the overall quantity of N that plants take up, the relationship between species'' abundance and available N in monoculture was largely driven by plant N uptake in this first year of growth.     

Utilisation of plant functional diversity in wildflower strips for the delivery of multiple agroecosystem services

Increased plant diversity in cropping systems can play an important role in agriculture by enhancing arthropod‐mediated ecosystem services, including biological control and pollination. However, there is limited research investigating the concurrent influence of plant functional diversity within cultivated systems on different arthropod functional groups, the provision of multiple ecosystem services, and crop yield. During a field experiment, repeated over 2 years, we measured the effect of increasing plant functional diversity on community structure of arthropod visitors, the abundance of multiple pests and induced crop damage, and fruit production in two varieties of tomato. Plant resources (floral and extra‐floral nectar and pollen) were included within experimental plots in four levels, with each level increasing the plant functional group richness, based on floral morphology and availability of resources, in a replacement series. The presence of sown flower mixtures in experimental plots was associated with increased abundance and diversity of natural enemy functional groups and an enhanced abundance of bees (Hymenoptera: Apiformes). However, we only detected relatively small variability in arthropod visitors among types of mixtures, and increased abundance of natural enemies did not translate into stronger pest suppression or reduced crop damage. Lepidoptera pest damage was significantly higher in plots adjacent to wildflower strips, an ecosystem disservice, but a significantly higher crop productivity was recorded from these plots. Our results provide evidence that inclusion of non‐crop plant resources in agroecosystems can improve the conservation of beneficial arthropods and may lead to increased crop productivity.     

农田生物多样性与害虫综合治理

在现代农田生态系统中 ,人类为了满足自身的需要 ,通常把自然界的植物群落改造成大面积种植单种特定的作物 ,人为地排除其他植物种类的竞争以提高作物的产量。由于单一化的作物不断取代自然植被 ,降低了农田的物种和生境多样性 ,结果导致农田生态系统的不稳定和害虫问题的更加恶化。影响农田生物多样性的因素很多 ,如地理位置、气候类型、环境条件、作物品种、种间关系、人类的栽培活动等。根据现有的生态学原理和研究成果 ,人类可以通过采用适当的措施来恢复和强化农田生物多样性 ,提高天敌的控制潜能 ,减少害虫发生的可能性 ;这些措施包括合理安排混作的时空格局 ,通过轮作进行间断性耕种 ,对多年生作物采用地面覆盖植被 ,利用不同品种以提高作物的遗传多样性 ,等等。在设计农田生物多样性的管理策略时 ,必须同时考虑当地气候、地理 ,植被 ,作物 ,土壤等因素的变化 ,因为在特定的生境条件下 ,这些因素可能增加或减少害虫发生的机会     

Mycorrhizal fungi as mediators of defence against insect pests in agricultural systems

• 1 Below‐ground organisms influence above‐ground interactions in both natural and agricultural ecosystems. Among the most important below‐ground organisms are mycorrhizal fungi, comprising ubiquitous and ancient plant mutualists that have significant effects on plant growth and fitness mediated by resource exchange with plants. In the present study, we focus on the effects of arbuscular mycorrhizal fungi (AMF) on crop defence against insect pests.
• 2 AMF alter the availability of resources used by crop plants to manufacture defences against pests and to compensate for pest damage. However, AMF also provide plants with nutrients that are known to increase insect performance. Through potentially opposing effects on plant nutritional quality and defence, mycorrhizal fungi can positively or negatively affect pest performance.
• 3 Additionally, AMF may directly affect gene expression and plant defence signalling pathways involved in the construction and induction of plant defences, and these effects are apparently independent of those caused by nutrient availability. In this way, AMF may still influence plant defences in the fertilized and highly managed systems typical of agribusiness.
• 4 Because AMF can affect plant tolerance to pest damage, they may have a significant impact on the shape of damage–yield relationships in crops. Potential mechanisms for this effect are suggested.
• 5 We highlight the need for continuing research on the effects of AMF identity and the abundance on crop defences and tolerance to pest attack. Much work is needed on the potential effects of mycorrhizal colonization on plant signalling and the induction of direct and indirect defences that may protect against pest damage.

     

Early growth and nutritional response to resource competition in cocoa-shade intercropped systems

Intercropping is often promoted for effective mutualism between species, thus compensating for external inputs. However, for optimal farm design resulting in superior production and nutrition, an accurate assessment of plant inter- and intra-specific competition is required. In predominant shade tree-cocoa (Theobroma cacao) systems, inconclusive evidence remains on species interactions, limitations to resource availability and subsequent growth and nutritional response, particularly in early growth. We examined cocoa biomass and foliar nutrition as well as nutrient supply through rates of decomposition and N mineralization after 1-year growth. Our approach employed fertilization and mixed planting treatments in an additive design of cocoa in monoculture (control), under artificial shade, and intercropped under two separate shade species (Terminalia superba and Newbouldia laevis). Intercropping had no effect on cocoa biomass production in comparison to monoculture cocoa. However, artificial shading stimulated foliage and root production both with and without fertilization, suggesting strong effects of light regulation on growth in the absence of belowground competition. Nutritionally, intercropping suppressed K uptake in cocoa foliage as K concentration was reduced by 20–25%, signifying dilution of this nutrient, presumably due to interspecific competition for mobile elements. Foliar N content under N. laevis was raised, where N concentration kept up with growth under this intercropped species. Intercropping also delayed decomposition rates, suggesting slower but sustained release of available nutrients into the topsoil. Cocoa under artificial shade, both with and without fertilization, exhibited the greatest nutrient responses as compared to unfertilized monoculture cocoa, where P uptake was stimulated most (175 and 112%), followed by K (69 and 71%), and then N (54 and 42%). Intercropping with shade trees failed to increase cocoa biomass, however, nutrient uptake was sustained for N and P, suggesting low interspecific competition. When fertilizers are undesirable or unavailable, intercropping of appropriately selected shade trees will not competitively suppress early growth of cocoa but will improve light regulation and nutritional status of cocoa saplings.     

Resource pulses of dead periodical cicadas increase the growth of American bellflower rosettes under competitive and non-competitive conditions

This experiment investigates how pulsed nutrient resources interact with intraspecific competition to influence biomass production and nutrient use efficiency in the American bellflower (Campanulastrum americanum). The competitive environment of these plants was manipulated by growing plants alone or with neighboring conspecifics, and the occurrence of pulsed nutrient resources was manipulated through the addition of 17-year periodical cicada (Magicicada septendecim) carcasses in order to simulate naturally occurring pulsed resources in this system. The addition of cicada carcasses increased mean plant biomass by 61 % compared to non-supplemented plants, while competition decreased mean plant biomass by 44 % compared to plants grown without competition; these effects were additive. In comparison, nutrient use efficiency decreased in fertilized plants (cicada-supplemented plants showed 20 % greater foliage nitrogen concentrations compared to non-supplemented plants), but was not significantly affected by the plant’s competitive environment. In addition, cicada supplementation did not significantly increase the biomass asymmetry in competing pairs of plants. These results suggest that these plants increased their total nutrient uptake at a timescale commensurate with the pulsed increase in nutrient availability due to cicada carcass decomposition.     

The role of belowground competition and plastic biomass allocation in altering plant mass–density relationships

Metabolic scaling theory (MST) predicts a ‘universal scaling law’ for plant mass–density relationships, but empirical observations are more variable. Possible explanations of this variability include plasticity in biomass allocation between the above‐ and belowground compartment and different modes of competition, which can be asymmetric or symmetric. Although complex interactions of these factors are likely to occur, so far the majority of modelling and empirical studies has focussed on mono‐factorial explanations. We here present a generic individual‐based model, which allows exploring the plant mass–density relationship in realistic settings by representing plasticity of biomass allocation and different modes of competition in the above‐ and belowground compartment. Plants grew according to an ontogenetic growth model derived from MST. To evaluate the behavior of the simulated plants related to the allocation patterns and to validate model predictions, we conducted greenhouse experiments with tree seedlings. The model reproduced empirical patterns both at the individual and population level. Without belowground resource limitation, aboveground processes dominated and the slopes of mass–density relationships followed the predictions of MST. In contrast, resource limitation led to an increased allocation of biomass to belowground parts of the plants. The subsequent dominance of symmetric belowground competition caused significantly shallower slopes of the mass–density relationship, even though the growth of individual plants followed MST. We conclude that changes in biomass allocation induced by belowground resource limitation explain the deviations from the mass–density relationship predicted by MST. Taking into account the plasticity of biomass allocation and its linkage to the above‐ and belowground competition is critical for fully representing plant communities, in particular for correctly predicting their response of carbon storage and sequestration to changing environmental conditions.     

A fungal endophyte of an annual weed reduces host competitive ability and confers associational protection to wheat

Epichloid fungal endophytes (Epichloë spp., Ascomycota: Clavicipitaceae) inhabit aerial tissues of several cool-season grasses, and enhance host growth and defence against herbivores. The presence of these symbionts can also affect interactions between the host and other non-epichloid plants. The role of an epichloid endophyte on interspecific competition has been tested using perennial grasses with contrasting results, but it has been scarcely tested using annual species in agroecosystems. We evaluated the impact of Epichloë-grass symbiosis on the competitive interaction between a non-host cereal crop (Triticum aestivum, wheat) and a host weed (Lolium multiflorum, ryegrass), growing in the presence of invertebrate herbivores (aphids) under no resource limitation. We conducted an outdoor mesocosm experiment with wheat plants growing in monoculture or in mixture with low or high proportions of ryegrass plants. Ryegrass plants presented either low (E-) or high (E+) incidence of Epichloë occultans (i.e. frequency of epichloid endophytic plants). We measured wheat vegetative and reproductive yield and its natural aphid infestation. Although epichloid endophyte incidence did not affect ryegrass biomass, wheat reproductive yield in mixtures (relative to wheat monocultures) was 45% higher when grown with E+ ryegrass plants than E- conspecific plants. Aphids preferred wheat plants grown with E- plants rather than wheat plants grown with E+ plants, but only in mixtures with high proportion of ryegrass. Our results demonstrate that epichloid endophyte incidence can decrease host competitive ability and confers associational protection to the non-endophytic neighbouring plants. Thus, ryegrass-endophyte symbiosis can increase crop yield by positive neighbourhood effects through different mechanisms probably related to the density of the weed. The benefits of this endosymbiont cannot be considered host-exclusive since they can be disseminated to non-endophytic plants. Furthermore, our results suggest that the epichloid endophyte incidence on annual weeds can contribute to agroecosystem sustainability by influencing pest management and increasing crop yield.     

Competition increasingly dominates the responsiveness of juvenile beech and spruce to elevated CO2 and/or O3 concentrations throughout two subsequent growing seasons

Saplings of Fagus sylvatica and Picea abies were grown in mono‐ and mixed cultures in a 2‐year phytotron study under all four combinations of ambient and elevated ozone (O₃) and carbon dioxide (CO₂) concentrations. The hypotheses tested were (1) that the competitiveness of beech rather than spruce is negatively affected by the exposure to enhanced O₃ concentrations, (2) spruce benefits from the increase of resource availability (elevated CO₂) in the mixed culture and (3) that the responsiveness of plants to CO₂ and O₃ depends on the type of competition (i.e. intra vs. interspecific). Beech displayed a competitive disadvantage when growing in mixture with spruce: after two growing seasons under interspecific competition, beech showed significant reductions in leaf gas exchange, biomass development and crown volume as compared with beech plants growing in monoculture. In competition with spruce, beech appeared to be nitrogen (N)‐limited, whereas spruce tended to benefit in terms of its plant N status. The responsiveness of the juvenile trees to the atmospheric treatments differed between species and was dominated by the type of competition: spruce growth benefited from elevated CO₂ concentrations, while beech growth suffered from the enhanced O₃ regime. In general, interspecific competition enhanced these atmospheric treatment effects, supporting our hypotheses. Significant differences in root : shoot biomass ratio between the type of competition under both elevated O₃ and CO₂ were not caused by readjustments of biomass partitioning, but were dependent on tree size. Our study stresses that competition is an important factor driving plant development, and suggests that the knowledge about responses of plants to elevated CO₂ and/or O₃, acquired from plants growing in monoculture, may not be transferred to plants grown under interspecific competition as typically found in the field.     

Counterbalancing effects of competition for resources and facilitation against grazing in alpine snowbed communities

Alpine snowbeds are habitats where the major limiting factors for plant growth are herbivory and a small time window for growth due to late snowmelt. Despite these limitations, snowbed vegetation usually forms a dense carpet of palatable plants due to favourable abiotic conditions for plant growth within the short growing season. These environmental characteristics make snowbeds particularly interesting to study the interplay of facilitation and competition. We hypothesised an interplay between resource competition and facilitation against herbivory. Further, we investigated whether these predicted neighbour effects were species‐specific and/or dependent on ontogeny, and whether the balance of positive and negative plant–plant interactions shifted along a snowmelt gradient. We determined the neighbour effects by means of neighbour removal experiments along the snowmelt gradient, and linear mixed model analyses. The results showed that the effects of neighbour removal were weak but generally consistent among species and snowmelt dates, and depended on whether biomass production or survival was considered. Higher total biomass and increased fruiting in removal plots indicated that plants competed for nutrients, water, and light, thereby supporting the hypothesis of prevailing competition for resources in snowbeds. However, the presence of neighbours reduced herbivory and thereby also facilitated survival. For plant growth the facilitative effects against herbivores in snowbeds counterbalanced competition for resources, leading to a weak negative net effect. Overall the neighbour effects were not species‐specific and did not change with snowmelt date. Our finding of counterbalancing effects of competition and facilitation within a plant community is of special theoretical value for species distribution models and can explain the success of models that give primary importance to abiotic factors and tend to overlook interrelations between biotic and abiotic effects on plants.     

Light partitioning in experimental grass communities

Through complementary use of canopy space in mixtures, aboveground niche separation has the potential to promote species coexistence and increase productivity of mixtures as compared to monocultures. We set up an experiment with five perennial grass species which differed in height and their ability to compete for light to test whether plants partition light under conditions where it is a limiting resource, and if this resource partitioning leads to increased biomass production in mixtures (using relative yield-based methods). Further, we present the first application of a new model of light competition in plant communities. We show that under conditions where biomass production was high and light a limiting resource, only a minority of mixtures outperformed monocultures and overyielding was slight. The observed overyielding could not be explained by species differences in canopy structure and height in monoculture and was also not related to changes in the canopy traits of species when grown in mixture rather than monoculture. However, where overyielding occurred, it was associated with higher biomass density and light interception. In the new model of competition for light, greater light use complementarity was related to increased total energy absorption. Future work should address whether greater canopy space-filling is a cause or consequence of overyielding.     

接种AMF对菌根植物和非菌根植物竞争的影响

为了研究丛枝菌根真菌(arbuscular mycorrhizal fungus, AMF)对菌根植物与非菌根植物种间竞争的影响,以玉米(菌根植物)和油菜(非菌根植物)作为供试植物,分别进行间作、尼龙网分隔和单作,模拟这两种植物之间不同的竞争状态,接种丛枝菌根真菌Glomus intraradices和Glomus mosseae,比较菌根植物和非菌根植物的生长和磷营养状况,分析AMF侵染对植物种间竞争作用的影响。结果显示,与单作相比,间作模式下玉米的生物量及磷营养状况有所降低,但其菌根依赖性却有所提高。与不接种相比,接种处理显著降低了间作体系油菜根系的磷含量和磷吸收量,但趋于改善菌根植物玉米的磷营养状况。因此,接种AMF可以降低非菌根植物的磷营养状况及生物量,使得菌根植物的相对竞争能力明显提高,说明AMF在维持物种多样性方面有着重要的作用。     

Neighbouring monocultures enhance the effect of intercropping on the turnip root fly (Delia floralis)

Knowledge of insect behaviour is essential for accurately interpreting studies of diversification and to develop diversified agroecosystems that have a reliable pest‐suppressive effect. In this study, we investigated the egg‐laying behaviour of the turnip root fly, Delia floralis (Fall.) (Diptera: Anthomyiidae), in an intercrop‐monoculture system. We examined both the main effect of intercropping and the effect on oviposition in the border zone between a cabbage monoculture [Brassica oleracea L. var. capitata (Brassicaceae)] and a cabbage‐red clover intercropping system [Trifolium pratense L. (Fabaceae)]. To investigate the border‐effect, oviposition was measured along a transect from the border between the treatments to the centre of experimental plots. Intercropping reduced the total egg‐laying of D. floralis with 42% in 2003 and 55% in 2004. In 2004, it was also found that the spatial distribution of eggs within the experimental plots was affected by distance from the adjoining treatment. The difference in egg‐laying between monoculture and intercropping was most pronounced close to the border, where egg‐laying was 68% lower on intercropped plants. This difference in egg numbers decreased gradually up to a distance of 3.5 m from the border, where intercropped plants had 43% fewer eggs than the corresponding monocultured plants. The reason behind this oviposition pattern is most likely that flies in intercropped plots have a higher probability of entering the monoculture if they are close to the border than if they are in the centre of a plot. When entering the monoculture, flies can pursue their egg‐laying behaviour without being disrupted by the clover. As the final decision to land is visually stimulated, flies could also be attracted to fly from the intercropped plots into the monoculture, where host plants are more visually apparent. Visual cues could also hinder flies in a monoculture from entering an intercropped plot. Other possible patterns of insect attack due to differences in insect behaviour are discussed, as well as the practical application of the results of this study.     

Responses of nectar‐feeding birds to floral resources at multiple spatial scales

The responses of animal pollinators to the spatially heterogeneous distribution of floral resources are important for plant reproduction, especially in species‐rich plant communities. We explore how responses of pollinators to floral resources varied across multiple spatial scales and studied the responses of two nectarivorous bird species (Cape sugarbird Promerops cafer, orange‐breasted sunbird Anthobaphes violacea) to resource distributions provided by communities of co‐flowering Protea species (Proteaceae) in South African fynbos. We used highly resolved maps of about 125 000 Protea plants at 27 sites and estimated the seasonal dynamics of standing crop of nectar sugar for each plant to describe the spatiotemporal distribution of floral resources. We recorded avian population sizes and the rates of bird visits to > 1300 focal plants to assess the responses of nectarivorous birds to floral resources at different spatial scales. The population sizes of the two bird species responded positively to the amount of sugar resources at the site scale. Within sites, the effects of floral resources on pollinator visits to plants varied across scales and depended on the resources provided by individual plants. At large scales (radii > 25 m around focal plants), high sugar density decreased per‐plant visitation rates, i.e. plants competed for animal pollinators. At small scales (radii < 5 m around focal plants), we observed either competition or facilitation for pollinators between plants, depending on the sugar amount offered by individual focal plants. In plants with copious sugar, per‐plant visitation rates increased with increasing local sugar density, but visitation rates decreased in plants with little sugar. Our study underlines the importance of scale‐dependent responses of pollinators to floral resources and reveals that pollinators’ responses depend on the interplay between individual floral resources and local resource neighbourhood.     

A negative heterogeneity–diversity relationship found in experimental grassland communities

Recent meta-analyses and simulation studies have suggested that the relationship between soil resource heterogeneity and plant diversity (heterogeneity–diversity relationship; HDR) may be negative when heterogeneity occurs at small spatial scales. To explore different mechanisms that can explain a negative HDR, we conducted a mesocosm experiment combining a gradient of soil nutrient availability (low, medium, high) and scale of heterogeneity (homogeneous, large-scale heterogeneous, small-scale heterogeneous). The two heterogeneous treatments were created using chessboard combinations of low and high fertility patches, and had the same overall fertility as the homogeneous medium treatment. Soil patches were designed to be relatively larger (156 cm²) and smaller (39 cm²) than plant root extent. We found plant diversity was significantly lower in the small-scale heterogeneous treatment compared to the homogeneous treatment of the same fertility. Additionally, low fertility patches in the small-scale heterogeneous treatment had lower diversity than patches of the same size in the low fertility treatment. Shoot and root biomass were larger in the small-scale heterogeneous treatment than in the homogeneous treatment of the same fertility. Further, we found that soil resource heterogeneity may reduce diversity indirectly by increasing shoot biomass, thereby enhancing asymmetric competition for light resources. When soil resource heterogeneity occurs at small spatial scales it can lower plant diversity by increasing asymmetric competition belowground, since plants with large root systems can forage among patches and exploit soil resources. Additionally, small-scale soil heterogeneity may lower diversity indirectly, through increasing light competition, when nutrient uptake by competitive species increases shoot biomass production.     

Towards an understanding of the mechanisms of tolerance: compensating for herbivore damage by enhancing a mutualism

Abstract.   1. Traditionally, losses in plant fitness or yield resulting from insect damage have been redressed by reducing pest populations using insecticides or biocontrol; these approaches rely on the untested assumption that reduced plant fitness or yield is caused by diminished resources available to damaged plants.
2. By experimentally manipulating pollination and damage levels independently, it is shown that pollination, as well as lack of resources, may be limiting to damaged plants in a model insect-pollinated crop, cantaloupe.
3. With enhanced pollination, damaged plants produce as much fruit as undamaged plants, even under high damage levels. In contrast, damaged plants without supplemental pollination produced significantly less fruit than undamaged plants.
4. This approach is unique in shifting the focus away from reducing pest populations and toward enhancing mutualistic interactions. It avoids risks posed by insecticides (which also kill pollinators) and by biocontrol agents, known threats to native species.
5. Determining the mechanism underlying compensation sheds light on recovery from insect damage in both natural and managed systems. These results have a bearing on managing native plant populations suffering from pollinator declines.
6. Finally, it may be predicted that resources could limit tolerance to herbivore damage in resource-poor or high competition environments, whereas pollination may limit tolerance when resource levels are high.     

Costs of Defense and a Test of the Carbon-Nutrient Balance and Growth-Differentiation Balance Hypotheses for Two Co-Occurring Classes of Plant Defense

One of the goals of chemical ecology is to assess costs of plant defenses. Intraspecific trade-offs between growth and defense are traditionally viewed in the context of the carbon-nutrient balance hypothesis (CNBH) and the growth-differentiation balance hypothesis (GDBH). Broadly, these hypotheses suggest that growth is limited by deficiencies in carbon or nitrogen while rates of photosynthesis remain unchanged, and the subsequent reduced growth results in the more abundant resource being invested in increased defense (mass-balance based allocation). The GDBH further predicts trade-offs in growth and defense should only be observed when resources are abundant. Most support for these hypotheses comes from work with phenolics. We examined trade-offs related to production of two classes of defenses, saponins (triterpenoids) and flavans (phenolics), in Pentaclethra macroloba (Fabaceae), an abundant tree in Costa Rican wet forests. We quantified physiological costs of plant defenses by measuring photosynthetic parameters (which are often assumed to be stable) in addition to biomass. Pentaclethra macroloba were grown in full sunlight or shade under three levels of nitrogen alone or with conspecific neighbors that could potentially alter nutrient availability via competition or facilitation. Biomass and photosynthesis were not affected by nitrogen or competition for seedlings in full sunlight, but they responded positively to nitrogen in shade-grown plants. The trade-off predicted by the GDBH between growth and metabolite production was only present between flavans and biomass in sun-grown plants (abundant resource conditions). Support was also only partial for the CNBH as flavans declined with nitrogen but saponins increased. This suggests saponin production should be considered in terms of detailed biosynthetic pathway models while phenolic production fits mass-balance based allocation models (such as the CNBH). Contrary to expectations based on the two defense hypotheses, trade-offs were found between defenses and photosynthesis, indicating that studies of plant defenses should include direct measures of physiological responses.