Growth pattern and carbon allocation to volatile leaf terpenes under nitrogen-limiting conditions in Heterotheca subaxillaris (Asteraceae)

Summary Rosettes of Heterotheca subaxillaris were grown at four levels of nitrate. Individual leaf volatile mono- and sesquiterpene content, leaf nitrogen content, and root and shoot dry weight were measured on individual leaves every two weeks for 18 weeks. Rosettes with the highest nitrate availability had 2.2-fold greater leaf nitrogen levels compared to plants with the lowest availability. As nitrate availability became increasingly limited, carbon allocation to both volatile leaf terpenes and root growht increased. Leaf mono- and sesquiterpene content was greatest in the young leaves of individuals growing at the lowest nitrate availability conditions. Higher levels of carbon-based herbivore-deterring chemicals in nitrate-limited plants may increase net productivity through retention of nitrogen that would otherwise be lost to herbivory.     

Tight Coupling Between Shoot Level Foliar N and P,Leaf Area,and Shoot Growth in Arctic Dwarf Shrubs Under Simulated Climate Change

Nutrient availability limits productivity of arctic ecosystems, and this constraint means that the amount of nitrogen (N) in plant canopies is an exceptionally strong predictor of vegetation productivity. However, climate change is predicted to increase nutrient availability leading to increases in carbon sequestration and shifts in community structure to more productive species. Despite tight coupling of productivity with canopy nutrients at the vegetation scale, it remains unknown how species/shoot level foliar nutrients couple to growth, or how climate change may influence foliar nutrients–productivity relationships to drive changes in ecosystem carbon gain and community structure. We investigated the influence of climate change on arctic plant growth relationships to shoot level foliar N and phosphorus (P) in three dominant subarctic dwarf shrubs using an 18-year warming and nutrient addition experiment. We found a tight coupling between total leaf N and P per shoot, leaf area and shoot extension. Furthermore, a steeper shoot length-leaf N relationship in deciduous species (Vaccinium myrtillus and Vaccinium uliginosum) under warming manipulations suggests a greater capacity for nitrogen to stimulate growth under warmer conditions in these species. This mechanism may help drive the considerable increases in deciduous shrub cover observed already in some arctic regions. Overall, our work provides the first evidence at the shoot level of tight coupling between foliar N and P, leaf area and growth i.e. consistent across species, and provides mechanistic insight into how interspecific differences in alleviation of nutrient limitation will alter community structure and primary productivity in a warmer Arctic.     

Herbivory-induced overcompensation and resource-dependent production of extrafloral nectaries in Luffa cylindrica (Cucurbitaceae)

Extrafloral nectaries (EFNs) are nectar secretory structures involved in the indirect defense of plants. In the sponge gourd (Luffa cylindrica), EFNs commonly occur on the lower surface of leaf blades and stipules and remain functional until leaf senescence. To test the hypothesis that the development of EFNs is influenced by herbivore damage and resource availability, we grew Luffa cylindrica under different concentrations of Hoagland's nutrient solution (nutrient-poor conditions: 10%, 50%; and control condition: 100%) and two herbivory treatments (damaged and undamaged leaves). We collected ten leaves from treated plants to quantify leaf area and EFN density. Overall, leaf area increased and EFN decreased in damaged plants, but this significantly depended on nutritional status. In undamaged plants, EFN density tended to remain constant, whereas foliar area increased with nutrient input. Under herbivory, foliar area increased at 10% but decreased at 50 and 100% of nutrients in relation to undamaged plants, whereas EFN density tended to increase with nutrient availability to exceed undamaged plants under control concentrations. Plants under nutrient-poor conditions subjected to herbivory exhibited an increased foliar area, characterizing a compensatory mechanism. Our results suggest that herbivore-induced indirect defense is a damage- and resource-dependent response in Luffa cylindrica. These findings contribute to understanding the factors that modulate indirect defenses and plant-herbivore-environment interactions.     

Human disturbance promotes herbivory by leaf‐cutting ants in the Caatinga dry forest

Anthropogenic disturbances are known to modify plant–animal interactions such as those involving the leaf‐cutting ants, the most voracious and proliferating herbivore across human‐modified landscapes in the Neotropics. Here, we evaluate the effect of chronic anthropogenic disturbance (e.g., firewood collection, livestock grazing) and vegetation seasonality on foraging area, foliage availability in the foraging area, leaf consumption and herbivory rate of the leaf‐cutting ant Atta opaciceps in the semiarid Caatinga, a mosaic of dry forest and scrub vegetation in northeast Brazil. Contrary to our initial expectation, the foraging area was not affected by either disturbance intensity or the interaction between season and disturbance intensity. However, leaf consumption and herbivory rate were higher in more disturbed areas. We also found a strong effect of seasonality, with higher leaf consumption and herbivory rate in the dry season. Our results suggest that the foraging ecology of leaf‐cutting ants is modulated by human disturbance and seasonality as these two drivers affect the spectrum and the amount of resources available for these ants in the Caatinga. Despite the low productivity of Caatinga vegetation, the annual rates of biomass consumption by A. opaciceps are similar to those reported from other leaf‐cutting ants in rain forests and savannas. This is made possible by maintaining high foraging activity even in the peak of the dry season and taking benefit from any resource available, including low‐quality items. Such compensation highlights the adaptive capacity of LCA to persist or even proliferate in human‐modified landscapes from dry to rain forests.     

Interannual changes in folivory and bird insectivory along a natural productivity gradient in northern Patagonian forests

Trophic regulation models suggest that the magnitude of herbivory and predation (top-down forces) should vary predictably with habitat productivity. Theory also indicates that temporal abiotic variation and within-trophic level heterogeneity both affect trophic dynamics, but few studies addressed how these factors interact over broad-scale environmental gradients. Here we document herbivory from leaf-feeding insects along a natural rainfall/productivity gradient in Nothofagus pumilio forests of northern Patagonia, Argentina, and evaluate the impact of insectivorous birds on foliar damage experienced by tree saplings at each end of the gradient. The study ran over three years (1997–2000) comprising a severe drought (1998–1999), which allowed us to test how climatic events alter top-down forces. Foliar damage tended to increase towards the xeric, least productive forests. However, we found a predictable change of insect guild prevalence across the forest gradient. Leaf miners accounted for the greater damage recorded in xeric sites, whereas leaf chewers dominated in the more humid and productive forests. Interannual folivory patterns depended strongly on the feeding guild and forest site. Whereas leaf-miner damage decreased during the drought in xeric sites, chewer damage increased after the drought in the wettest site. Excluding birds did not affect leaf damage from miners, but generally increased chewer herbivory on hydric and xeric forest saplings. Indirect effects elicited by bird exclusion became most significant after the drought, when total folivory levels were higher. Thus, interannual abiotic heterogeneity markedly influenced the amount of folivory and strength of top-down control observed across the forest gradient. Moreover, our results suggest that spatial turnovers between major feeding guilds may need be considered to predict the dynamics of insect herbivory along environmental gradients.     

Altitudinal patterns in host suitability for forest insects

Conspecific trees growing at high and low-elevations encounter different growing conditions and may vary in their suitability as hosts for herbivorous insects. Mountain tree populations may be more resistant to herbivory if low temperatures constrain growth more than they constrain photosynthesis, resulting in increased secondary metabolism (temperature hypothesis). Alternatively, mountain trees may be fertilized by atmospheric nitrogen deposition and become more palatable to insects (atmospheric deposition hypothesis). We evaluated these two hypotheses by comparing high- and low-elevation trees with insect bioassays and analyses of foliar nitrogen and condensed tannin. Contrary to the temperature hypothesis, high-elevation foliage had higher leaf nitrogen (six of six tree species) and allowed higher growth rates of Lymantria dispar larvae (five of six tree species). The nitrogen deposition hypothesis was broadly supported by measurements from two mountains showing that high-elevation trees tended to have higher leaf nitrogen, lower leaf tannins, and support higher insect growth performance than conspecific trees from lower elevations. The deposition hypothesis was further supported by fertilization studies showing that simulated atmospheric nitrogen deposition changed the foliar chemistry of valley trees to resemble that of high-elevation trees. Predictions that the altitudinal gradient in foliar chemistry and host suitability should be steepest on mountains receiving more deposition were largely not supported, but interpretations are complicated by lack of replication among mountains. In the northeastern United States, increased host suitability of high-elevation trees seems sufficient to influence the population dynamics and community composition of herbivores. Atmospheric nitrogen deposition offers a promising hypothesis to explain and predict some important spatial patterns in herbivory. Received: 21 September 1997 / Accepted: 12 June 1998     

Seasonal patterns of herbivory,leaf traits and productivity consumption in dry and wet Patagonian forests

1. Endemic herbivory can influence forest ecosystem function, but how annual productivity consumption relates to seasonal resource utilisation by folivore guilds remains poorly understood. 2. Monthly changes in leaf damage and foliage traits were monitored in ‘dry’ and ‘wet’Nothofagus pumilio (Fagales: Nothofagaceae) deciduous forests in northern Patagonia, Argentina. Herbivore‐induced leaf abscission was assessed and foliar productivity consumption was measured in the canopy and in litterfall harvests. 3. Seasonal damage ranged from 8% to 32% in dry forest, but remained below 5% in wet forest although foliar quality was higher in the latter. In dry forest, dominant guilds were temporally separated; leaf miners consumed younger foliage in spring to early summer, whereas leaf tiers prevailed in late summer to autumn. In wet forest, damage created by external chewers was concentrated in early summer. 4. Insect damage induced premature leaf abscission, especially in dry forest. Although foliar production in wet forest doubled that in dry forest, the percentage of productivity lost to folivores was higher in dry (14–20%) than in wet (1.2–1.8%) forest. 5. The overall greater impact of herbivory in dry forest canopies countered the expectation that consumption would increase with plant productivity and nutritional quality. Lower temperatures and a shorter growing season are likely to constrain folivory in wet forest stands.     

Geographic patterns of plant–herbivore interactions are driven by soil fertility

Aims Geographic patterns of the intensity of plant herbivory in relation to climate factors have garnered little general support and appear to be species specific. However, plant–herbivore interactions are also driven by resource availability, such as soil nutrient content, and it remains unclear whether broad-scale variation in soil factors is reflected in herbivore consumption rates across species’ ranges. Additionally, we know little of how intraspecific variation in tissue quality associates with edaphic and climatic factors, and how this variation controls herbivore consumption. The resource availability hypothesis (RAH) predicts that plant individuals growing in low-resource environments will have lower leaf nutritional quality and more constitutive defenses, which will result in lower rates of leaf consumption.     

Are gas exchange responses to resource limitation and defoliation linked to source:sink relationships?

Productivity of trees can be affected by limitations in resources such as water and nutrients, and herbivory. However, there is little understanding of their interactive effects on carbon uptake and growth. We hypothesized that: (1) in the absence of defoliation, photosynthetic rate and leaf respiration would be governed by limiting resource(s) and their impact on sink limitation; (2) photosynthetic responses to defoliation would be a consequence of changing source:sink relationships and increased availability of limiting resources; and (3) photosynthesis and leaf respiration would be adjusted in response to limiting resources and defoliation so that growth could be maintained. We tested these hypotheses by examining how leaf photosynthetic processes, respiration, carbohydrate concentrations and growth rates of Eucalyptus globulus were influenced by high or low water and nitrogen (N) availability, and/or defoliation. Photosynthesis of saplings grown with low water was primarily sink limited, whereas photosynthetic responses of saplings grown with low N were suggestive of source limitation. Defoliation resulted in source limitation. Net photosynthetic responses to defoliation were linked to the degree of resource availability, with the largest responses measured in treatments where saplings were ultimately source rather than sink limited. There was good evidence of acclimation to stress, enabling higher rates of C uptake than might otherwise have occurred.     

A Temporally Explicit Production Efficiency Model for Fuel Load Allocation in Southern Africa

Abstract We present a regional fuel load model (1 km² spatial resolution) applied in the southern African savanna region. The model is based on a patch-scale production efficiency model (PEM) scaled up to the regional level using empirical relationships between patch-scale behavior and multi-source remote sensing data (spatio-temporal variability of vegetation and climatic variables). The model requires the spatial distribution of woody vegetation cover, which is used to determine separate respiration rates for tree and grass. Net primary production, grass and tree leaf death, and herbivory are also taken into account in this mechanistic modeling approach. The fuel load model has been calibrated and validated from independent measurements taken from savanna vegetation in Africa southward from the equator. A sensitivity analysis on the effect of climate variables (incoming radiation, air temperature, and precipitation) has been conducted to demonstrate the strong role that water availability has in determining productivity and subsequent fuel load over the southern African region. The model performance has been tested in four different areas representative of a regional increasing rainfall gradient—Etosha National Park, Namibia, Mongu and Kasama, Zambia, as well as in Kruger National Park, South Africa. Within each area, we analyze model output from three different magnitudes of canopy coverage (<5, 30, and 50%). We find that fuel load ranges predicted by the model are globally in agreement with field measurements for the same year. High rainfall sustains green herbaceous production late in the dry season and delays tree leaf litter production. Effect of water on production varies across the rainfall gradient with delayed start of green material production in more arid regions.     

Invertebrate herbivory increases along an experimental gradient of grassland plant diversity

Plant diversity is a key driver of ecosystem functioning best documented for its influence on plant productivity. The strength and direction of plant diversity effects on species interactions across trophic levels are less clear. For example, with respect to the interactions between herbivorous invertebrates and plants, a number of competing hypotheses have been proposed that predict either increasing or decreasing community herbivory with increasing plant species richness. We investigated foliar herbivory rates and consumed leaf biomass along an experimental grassland plant diversity gradient in year eight after establishment. The gradient ranged from one to 60 plant species and manipulated also functional group richness (from one to four functional groups—legumes, grasses, small herbs, and tall herbs) and plant community composition. Measurements in monocultures of each plant species showed that functional groups differed in the quantity and quality of herbivory damage they experienced, with legumes being more damaged than grasses or non-legume herbs. In mixed plant communities, herbivory increased with plant diversity and the presence of two key plant functional groups in mixtures had a positive (legumes) and a negative (grasses) effect on levels of herbivory. Further, plant community biomass had a strong positive impact on consumed leaf biomass, but little effect on herbivory rates. Our results contribute detailed data from a well-established biodiversity experiment to a growing body of evidence suggesting that an increase of herbivory with increasing plant diversity is the rule rather than an exception. Considering documented effects of herbivory on other ecosystem functions and the increase of herbivory with plant diversity, levels of herbivory damage might not only be a result, but also a trigger within the diversity–productivity relationship.     

自然植被净第一性生产力模型初探

本文根据植物的生理生态学特点及联系能量平衡方程和水量平衡方程的区域蒸散模式建立了联系植物生理生态学特点和水热平衡关系的植物的净第一性生产力模型：该模型的建立为宏观地确定地带性景观的生产潜力、植物净第一性生产力的区域分布和全球分布,以及全球变化的影响提供了理论基础,对于合理地利用气候资源,扬长避短,充分发挥气候生产潜力,最大限度地提高植物的产量具有重要的意义。     

Environmental influences on growth and defence responses of the invasive shrub,Lonicera maackii,to simulated and real herbivory in the juvenile stage

Background and Aims

Tolerance and defence against herbivory are among the many mechanisms attributed to the success of invasive plants in their novel ranges. Because tolerance and defence against herbivory differ with the ontogeny of a plant, the effects of herbivore damage on plant fitness vary with ontogenetic stage and are compounded throughout a plant''s lifetime. Environmental stresses such as light and nutrient limitations can further influence the response of the plant. Much is known about the response of plants in the seedling and reproductive adult stages, but less attention has been given to the pre-reproductive juvenile stage.

Methods

Juvenile plants of the North American invasive Lonicera maackii were exposed to simulated herbivory under high and low light and nitrogen availability and growth, allocation patterns and foliar defensive chemistry were measured. In a second experiment, complete nutrient availability and damage type (generalist caterpillar or simulated) were manipulated.

Key Results

Juvenile plants receiving 50 % defoliation had lower total biomass and a higher root^:^shoot ratio than controls for all treatment combinations except low nitrogen/low light. Low light and defoliation increased root^:^shoot ratio. Light, fertilization and defoliation had little impact on foliar defensive chemistry. In the second experiment, there was a reduction in total biomass when caterpillar damage was applied. The root^:^shoot ratio increased under low soil fertility and was not affected by defoliation. Stem-diameter growth rates and specific leaf area did not vary by damage type or fertilization. Foliar protein increased through time, and more strongly in defoliated plants than in controls, while peroxidase activity and total flavonoids decreased with time. Overall, resource limitations were more influential than damage in the growth of juvenile L. maackii plants.

Conclusions

The findings illustrate that even when resources are limited, the tolerance and defence against herbivory of a woody invasive plant in the juvenile stage may contribute to the establishment and persistence of some species in a variety of habitats.     

Biotic and Abiotic Controls Over Canopy Function and Structure in Humid Hawaiian Forests

Foliar nitrogen (N) plays a key role in ecosystem function and dynamics, including processes such as photosynthesis, productivity, and decomposition. Aboveground carbon density (ACD Mg C ha^?1) represents a cumulative functional outcome of these and other ecosystem processes and is an important metric for monitoring current carbon stocks. Despite their importance, multiple interacting controls over landscape-level variation in foliar N and ACD are poorly understood. We assessed the relative importance of individual ecologically important state factors (climate, substrate, age, vegetation, and topography) associated with canopy foliar N and ACD throughout a humid forest landscape. We combined high-resolution remotely sensed data, machine learning, and field data to map and assess canopy foliar N and ACD patterns across a 5016-ha forest reserve in Hawai‘i. Distance to non-native forests had the largest relative influence on canopy foliar N concentration, followed by mean annual temperature (MAT), vegetation type, precipitation, soil, canopy height, and substrate age. In contrast, soil type was the strongest determinant of spatial variability in ACD, followed by precipitation, MAT, and vegetation type. Similar to foliar N, climate and vegetation variables were associated with ACD. However, soil type was found to be much more important in the ACD model (30%) than in the foliar N model (4%). Landscape-scale patterns in canopy foliar N and ACD are the result of shifts in vegetation type and composition, most likely due to species’ responses to past disturbances, current climate conditions, and available nutrients. Degradation of native forests and future climate changes could result in highly altered biogeochemical cycles.     

Interactive controls of herbivory and fluvial dynamics on landscape vegetation patterns on the Tanana River floodplain, interior Alaska

Aim  We examined the interactive effects of mammalian herbivory and fluvial dynamics on vegetation dynamics and composition along the Tanana River in interior Alaska.
Location  Model parameters were obtained from field studies along the Tanana River, Alaska between Fairbanks (64°50.50' N, 147°43.30' W) and Manley Hot Springs (65°0.0' N, 150°36.0' W).
Methods  We used a spatially explicit model of landscape dynamics (ALFRESCO) to simulate vegetation changes on a 1-year time-step. The model was run for 250 years and was replicated 100 times.
Results  Increases in herbivory decreased the proportion of early successional vegetation and increased the proportion of late successional vegetation on the simulated landscape. Erosion and accretion worked as antagonists to herbivory, increasing the amount of early successional vegetation and decreasing the amount of late successional vegetation. However, the interactive effects of herbivory and erosion/accretion were especially important in determining system response, particularly in early seral vegetation types. High erosion rates, when coupled with low herbivory, greatly increased the proportion of willow on the landscape. When coupled with high herbivory, however, they greatly increased the proportion of alder on the landscape. At low levels of herbivory, alder abundance peaked at intermediate levels of erosion/accretion.
Main conclusions  Neither erosion/accretion nor herbivory produced consistent landscape patterns that could be predicted independently of the other. These findings underscore the importance of the interactive effects of biotic and abiotic disturbances in shaping large-scale landscape vegetation patterns in boreal floodplain ecosystems – systems traditionally thought to be driven primarily by abiotic disturbance alone.     

Phytophagous insects enhance nitrogen flux in a desert creosotebush community

Summary We tested the hypothesis that herbivorous insects on desert shrubs contribute to short-term nitrogen cycling, and increase rates of nitrogen flux from nutrient rich plants. Creosotebush (Larrea tridentata) shrubs were treated with different combinations of fertilizer and water augmentations, resulting in different levels of foliage production and foliar nitrogen contents. Foliage arthropod populations, and nitrogen in canopy dry throughfall, wet throughfall and stemflow were measured to assess nitrogen flux rates relative to arthropod abundances on manipulated and unmanipulated shrubs over a one-month period during peak productivity. Numbers and biomass of foliage arthropods were significantly higher on fertilized shrubs. Sap-sucking phytophagous insects accounted for the greatest numbers of foliage arthropods, but leaf-chewing phytophagous insects represented the greatest biomass of foliage arthropods. Measured amounts of bulk frass (from leaf-chewing insects) were not significantly different among the various treatments. Amounts of nitrogen from dry and wet throughfall and stemflow were significantly greater under fertilized shrubs due to fine frass input from sap-sucking insects. Increased numbers and biomass of phytophagous insects on fertilized shrubs increased canopy to soil nitrogen flux due to increased levels of herbivory and excrement. Nitrogen excreted by foliage arthropods accounted for about 20% of the total one month canopy to soil nitrogen flux, while leaf litter accounted for about 80%.     

Tight coupling between leaf area index and foliage N content in arctic plant communities

The large spatial heterogeneity of arctic landscapes complicates efforts to quantify key processes of these ecosystems, for example productivity, at the landscape level. Robust relationships that help to simplify and explain observed patterns, are thus powerful tools for understanding and predicting vegetation distribution and dynamics. Here we present the same linear relationship between Leaf area index (LAI) and Total foliar nitrogen (TFN), the two factors determining the photosynthetic capacity of vegetation, across a wide range of tundra vegetation types in both northern Sweden and Alaska between leaf area indices of 0 and 1 m² m^–2, which is essentially the entire range of leaf area index values for the Arctic as a whole. Surprisingly, this simple relationship arises as an emergent property at the plant community level, whereas at the species level a large variability in leaf traits exists. As the relationship between LAI and TFN exists among such varied ecosystems, the arctic environment must impose tight constraints on vegetation canopy development. This relationship simplifies the quantification of vegetation productivity of arctic vegetation types as the two most important drivers of productivity can be estimated reliably from remotely sensed NDVI images.     

Effects of neighbouring vegetation on eucalypt seedlings at two sites subject to different levels of abiotic stress

Abstract In spite of numerous studies on the effect of nutrient levels and/or standing crop on the intensity of resource competition the debate has not been resolved. Field studies that have used natural productivity gradients have generally supported the argument that competitive intensity and resource availability are positively correlated, whereas studies that have used artificial resource gradients have generally refuted the same argument. Here we report the results from study in which both approaches were used within the same system. We studied two species of eucalypt that occupy contrasting parts of the same landscape: Eucalyptus camaldulensis, found mostly along creek lines and in valleys with deep alluvial soils, and Eucalyptus microcarpa, found on hillsides and ridges with shallow soils. We studied the response of seedlings of the two species to the combined effects of competition and manipulated nutrient levels in a glasshouse experiment, and also investigated their responses to removal of neighbouring plants in the field. Eucalyptus microcarpa was less responsive to increased resource availability, which is consistent with one of the principal assumptions of Grime’s C‐S‐R model. In the glasshouse experiment both species of eucalypt responded in a qualitatively similar fashion to the combined effects of resource availability and competition: release from competition resulted in increased growth, but only in pots that received additional resources. In the field we found that neighbouring vegetation could severely limit the establishment of E. camaldulensis but the removal of neighbouring vegetation did not affect the performance of E. microcarpa seedlings. Eucalyptus camaldulensis seedlings suffered high levels of damage from herbivores. Our results thus generally support the predications of the C‐S‐R model, however, they indicate that the effects of competition and herbivory may be heavily confounded.     

Effects of plant quality and ant defence on herbivory rates in a neotropical ant‐plant

1. Understanding the degree to which populations and communities are limited by both bottom‐up and top‐down effects is still a major challenge for ecologists, and manipulation of plant quality, for example, can alter herbivory rates in plants. In addition, biotic defence by ants can directly influence the populations of herbivores, as demonstrated by increased rates of herbivory or increased herbivore density after ant exclusion. The aim of this study was to evaluate bottom‐up and top‐down effects on herbivory rates in a mutualistic ant‐plant. 2. In this study, the role of Azteca alfari ants as biotic defence in individuals of Cecropia pachystachya was investigated experimentally with a simultaneous manipulation of both bottom‐up (fertilisation) and top‐down (ant exclusion) factors. Four treatments were used in a fully factorial design, with 15 replicates for each treatment: (i) control plants, without manipulation; (ii) fertilised plants, ants not manipulated; (iii) unfertilised plants and excluded ants and (iv) fertilised plants and ants excluded. 3. Fertilisation increased the availability of foliar nitrogen in C. pachystachya, and herbivory rates by chewing insects were significantly higher in fertilised plants with ants excluded. 4. Herbivory, however, was more influenced by bottom‐up effects – such as the quality of the host plant – than by top‐down effects caused by ants as biotic defences, reinforcing the crucial role of leaf nutritional quality for herbivory levels experienced by plants. Conditionality in ant defence under increased nutritional quality of leaves through fertilisation might explain increased levels of herbivory in plants with higher leaf nitrogen.     

Effects of genotype and nutrient availability on phytochemistry of trembling aspen (Populus tremuloides Michx.) during leaf senescence

We documented temporal patterns in phytochemical composition of Populus tremuloides Michx. during leaf senescence, and the influence of genotype and soil nutrient availability on such patterns. Levels of foliar nitrogen, carbohydrates, phenolic glycosides and condensed tannins were quantified for four aspen genotypes grown in a common garden, with low and high levels of soil nutrients. Levels of all compounds tended to decline over time, although the magnitude of change was influenced by plant genotype and nutrient availability. Genetic variation in concentrations of phytochemicals was much greater for phenolic glycosides and tannins than for nitrogen and carbohydrates, and these phenolic signatures generally persisted through leaf abscission. Our results suggest that genotypic and nutrient effects on patterns of chemical change during senescence will likely influence the performance of late-season herbivores on aspen. Moreover, nutrient and especially genotypic variation in phytochemistry of abscised leaves is likely to affect litter decomposition rates.