Delayed induced changes in the biochemical composition of host plant leaves during an insect outbreak

In birch, Betula pubescens, herbivore-induced delayed induced resistance (DIR) of defoliated trees may cause a strong reduction in the potential fecundity of a geometrid folivore Epirrita autumnata. In this study, we examined the biochemical basis of DIR in birch leaves during a natural outbreak of E. autumnata. A set of experimental trees was defoliated at four sites by wild larvae in the peak year of the outbreak, whereas control trees were protected from defoliation by spraying with an insecticide. The biochemical composition of leaves was analysed in the following year and, although the DIR response was weak during this outbreak, causing less than a 20% reduction in the potential fecundity of E. autumnata, some consistent relationships between defoliation, biochemistry and pupal mass of E. autumnata suggested a general biochemical basis for the defoliation-induced responses in birch leaves. Total concentrations of nitrogen, sugars and acetone-insoluble residue (e.g. cell wall polysaccharides, cell-wall-bound phenolics, protein, starch, lignin and hemicellulose) were consistently lower, and total concentrations of phenolics, especially of gallotannins and soluble proanthocyanidins, were higher in the leaves of trees defoliated in the previous year than in those protected from defoliation. The capacity of tannins to precipitate proteins correlated with contents of gallotannins, and was highest in defoliated trees. The pupal mass of E. autumnata showed a strong, positive correlation with concentrations of nitrogen and sugars, and a negative correlation with the acetone-insoluble residue and gallotannins in foliage. Correlations with other measured biochemical traits were weak. The correlation coefficients between biochemical traits and pupal mass consistently had similar signs for both defoliated and insecticide–sprayed trees, suggesting that variation in leaf quality due to defoliation in the previous year was based on similar biochemical traits as variation for other reasons. We suggest that DIR is associated with reduced growth activity of leaves, and may be seen as a delay in the biochemical maturation of leaves in defoliated trees. This explains the high concentration of gallotannins in defoliated trees, a characteristic feature of young leaves. However, the lower content of nitrogen and the higher content of soluble proanthocyanidins in defoliated trees are traits usually characterising mature, not young, leaves, indicating defoliation-induced changes in chemistry in addition to modified leaf age. Our results emphasise the importance of understanding the natural changes in chemistry during leaf maturation when interpreting defoliation-induced changes in leaf biochemistry. Received: 26 January 1998 / Accepted: 10 April 1998     

Covariation of fluctuating asymmetry, herbivory and chemistry during birch leaf expansion

Fluctuating asymmetry (FA) is used to describe developmental instability in bilateral structures. In trees, high FA of leaves has been assumed to indicate the level of environmental or genetic stress, and for herbivores leaves from such trees have been shown to be in some cases (though not invariably) of higher quality compared to trees with symmetrical leaves. We demonstrated that FA of birch leaves correlated positively with growth rate of leaves, and with the amount of leaf biomass consumed by larvae of the geometrid Epirrita autumnata. Since asymmetry per se cannot define leaf quality for a herbivore, we determined the biochemical compounds which covary with the degree of foliage FA, in order to elucidate relationships between leaf FA, chemistry and herbivory. High foliar FA was characteristic of birches with high initial concentrations, and rapid seasonal decline in the concentrations of gallic acid and hydrolysable tannins, and with rapid seasonal changes in the concentrations of flavonoid-glycosides and sugars. In contrast, leaf FA was not related to concentrations of proanthocyanidins, protein-bound amino acids or soluble phenylalanine, the precursor of proanthocyanidins and proteins with aromatic amino acids. The positive correlation between leaf FA and consumption by E. autumnata was presumably related to the previously demonstrated compensatory consumption of E. autumnata to high concentrations of foliar gallotannins. Furthermore, sugars are well-known feeding stimulants. We propose that the variable results in studies correlating leaf FA and herbivory may stem from variable chemical associations of FA in different plants and of species-specific effects of compounds on insects. Received: 15 July 1999 / Accepted: 24 September 1999     

Effects of host shading on consumption and growth of the geometrid Epirrita autumnata: interactive roles of water, primary and secondary compounds

Shading is assumed to reduce allocation to plant phenolics and to defense in general. We here report the results of experimental shading of individual branches or whole canopies in mountain birch on foliar chemistry and on the growth and consumption of a geometrid, Epirritaautumnata. Branch‐wide shading tended to have at least as strong effects on both leaf chemistry and herbivore performance as canopy‐wide shading, indicating local responses of the host to shading. Responses to shading varied among the key leaf traits. Leaf water content was higher and toughness lower in shaded than in non‐shaded leaves. Leaf sugars were lower and protein‐bound and free amino acids higher in shaded than in control leaves. Sucrose and galactose were at high levels in unshaded branches adjacent to shaded ones, suggesting that partial shading enhanced translocation of sugars within canopies. Total phenolics and soluble proanthocyanidins were low in both shading treatments. Of the other phenolic groups, concentrations of gallotannins and cell‐wall‐bound proanthocyanidins did not differ between shaded and non‐shaded leaves. Epirrita larvae grew better in both types of shading treatments compared to either unshaded control trees or to unshaded branches in the branch‐shading trees. By far the most important correlate of larval growth was the amount of water consumed with leaf mass (r=0.94). When variance in water intake was standardized (also largely eliminating parallel variation in proteins), fructose and glucose still had significant positive correlations and proanthocyanidins negative with larval growth on control but not on shade leaves. Concentrations of several phenolic compounds correlated negatively with intake of dry matter and especially water, and different phenolics were important in shaded (gallotannins) and in control (flavonoids) leaves. Our findings strongly suggest that the effects of putatively defensive leaf traits on insect consumption and growth interact with nutritive leaf traits, particularly with water.     

Intraspecific variation in Eucalyptus secondary metabolites determines food intake by folivorous marsupials

Traditional approaches to the question of the effects of plant secondary metabolites on the feeding choices of folivores of Eucalyptus have focused on the tree species level, although numerous field studies of foraging behaviour have identified selection at the level of the individual trees. Attempts to relate these decisions to deterrency resulting from secondary leaf chemistry have been inconclusive because assays used have focused on broad groups of compounds such as “total” phenolics. In this study we have conducted no-choice feeding trials with two arboreal mammalian folivores, the common ringtail possum (Pseudocheirus peregrinus) and the koala (Phascolarctos cinereus), to measure deterrency of individual trees of two species of Eucalyptus, E. ovata and E. viminalis. Average daily intakes of E. ovata foliage by common ringtail possums ranged from 2.5 to 50 g kg^−0.75 body mass. Koala intakes of foliage from the same individual trees ranged from 22.4 to 36.3 g kg^−0.75 body mass. When fed foliage from different individual E. viminalis trees, common ringtail possums ate between 1.26 and 6.28 g kg^−0.75 body mass while koalas ate from 14.3 to 45.9 g kg^−0.75 body mass. Correlative analyses showed no relationships between feeding and several measures of nutritional quality, nor with total phenolics or condensed tannins. They did, however, identify two groups of plant secondary metabolites that may cause deterrency: terpenes, and a defined group of phenolic compounds, the diformylphloroglucinols (DFPs). Further bioassay experiments with common ringtail possums showed that only the DFPs could cause the effects seen with the foliage experiments at concentrations similar to those found in the leaves. We argue that, when in sufficiently high concentrations, DFPs determine the level of food intake by these animals irrespective of other questions of nutritional quality of the leaves. Received: 20 October 1997 / Accepted: 23 March 1998     

A High Concentration of Glucogallin, the Common Precursor of Hydrolyzable Tannins, Does Not Deter Herbivores

The diversity of structures of plant phenolic compounds suggests that their interactions with insect herbivores may be compound specific. In this study, we modified the natural covariances observed in mature leaves of mountain birch, Betula pubescens ssp. czerepanovii (Orlova) Hämet-Ahti, by supplying gallic acid, the common precursor of gallotannins, through the stems of cut branches. Only one gallotannin, glucogallin, was consequently increased, and responses to this change on larvae of Epirrita autumnata Bkh. were evaluated by choice and nonchoice experiments. Glucogallin-increased leaves were consumed equally to control leaves in a nonchoice situation and they were preferred by E. autumnata larvae when they had to choose. No other short-term postingestive effects in E. autumnata larvae were observed and therefore our studies did not suggest a defensive role for glucogallin.     

Nitrogen Controls on Fine Root Substrate Quality in Temperate Forest Ecosystems

Nitrogen controls on fine root substrate quality (that is, nitrogen and carbon-fraction concentrations) were assessed using nitrogen availability gradients in the Harvard Forest chronic nitrogen addition plots, University of Wisconsin Arboretum, Blackhawk Island, Wisconsin, and New England spruce-fir transect. The 27 study sites encompassed within these four areas collectively represented a wide range of nitrogen availability (both quantity and form), soil types, species composition, aboveground net primary production, and climatic regimes. Changes in fine root substrate quality among sites were most frequently and strongly correlated with nitrate availability. For the combined data set, fine root nitrogen concentration increased (adjusted R ² = 0.46, P < 0.0001) with increasing site nitrate availability. Fine root “extractive” carbon-fraction concentrations decreased (adjusted R ² = 0.32, P < 0.0002), “acid-soluble” compounds increased (adjusted R ² = 0.35, P < 0.0001), and the “acid-insoluble” carbon fraction remained relatively high and stable (combined mean of 48.7 ± 3.1% for all sites) with increasing nitrate availability. Consequently, the ratio of acid-insoluble C–total N decreased (adjusted R ² = 0.40, P < 0.0001) along gradients of increasing nitrate availability. The coefficients of determination for significant linear regressions between site nitrate availability and fine root nitrogen and carbon-fraction concentrations were generally higher for sites within each of the four study areas. Within individual study sites, tissue substrate quality varied between roots in different soil horizons and between roots of different size classes. However, the temporal variation of fine root substrate quality indices within specific horizons was relatively low. The results of this study indicate that fine root substrate quality increases with increasing nitrogen availability and thus supports the substrate quality component of a hypothesized conceptual model of nitrogen controls on fine root dynamics that maintains that fine root production, mortality, substrate quality, and decomposition increase with nitrogen availability in forest ecosystems in a manner that is analogous to foliage.     

Within-tree variability of mountain birch leaves causes variation in performance for Epirrita autumnata larvae

Within-tree variability in leaf characteristics of the mountain birch (Betula pubescens ssp. tortuosa) was bioassayed for the autumnal moth (Epirrita autumnata) by rearing larvae on birch leaves in a laboratory and measuring their growth, consumption rate, approximate digestibility, efficiency of conversion of ingested food and efficiency of conversion of digested food. Only short shoot leaves, i.e. leaves of the same age, were used. The highest hierarchical level, which included trees and ramets within trees, accounted for most of the total variance in almost all the measured traits. Short shoots (within branches) accounted for more of the variance than branches (within trees/ramets) in most of the traits. The results suggest that differences in leaf quality were reflected in larval growth mainly by differences in food utilization efficiencies (postingestive effects) and less by differences in consumption rate (preingestive effects). The observed within-tree variation is probably a consequence of the modular structure, sectoriality and partial functional independence of tree parts.     

Influence of leaf traits on the spatial distribution of insect herbivores associated with an overstorey rainforest tree

Summary The spatial distribution of insect herbivores associated with the Australian rainforest treeArgyrodendron actinophyllum (Sterculiaceae) was investigated by restricted canopy fogging. The foliage of this species was low in nitrogen and water content, and high in fibre content. Herbivore abundance was positively correlated with the amount of young foliage present within the samples and in adjacent samples, and with the nitrogen content of young leaves. In particular, the occurrence of phloem-feeders was correlated with the magnitude of translocation within the samples. The influence of leaf water content upon herbivore distribution was marginal, presumably because this factor is not limiting in rain-forest environments during the wet season, which usually coincides with the season of leaf-flush. Specific leaf weight, leaf size and foliage compactness had little or no apparent effect on herbivore distribution. Since the magnitude of leaf turnover affected both the quantity and the quality, as exemplified by translocation effects, of young foliage available, this factor may be critical to herbivores associated with evergreen rainforest trees which are particularly low in foliar nutrients, such asA. actinophyllum.     

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     

Oviposition preference and neonate performance of Mnesampela privata in relation to heterophylly in Eucalyptus dunnii and E. globulus

Abstract

• 1 The autumn gum moth, Mnesampela privata, is an endemic Australian geometrid that utilizes a number of species within the genus Eucalyptus as hosts. Based on field observations, the moth is thought to be leaf‐type specific for juvenile as opposed to adult eucalypt foliage.
• 2 Laboratory binary choice assays of the oviposition preference of host novice M. privata confirmed that eggs were more likely to be laid upon juvenile rather than adult foliage of Eucalyptus dunnii and two subspecies of E. globulus. This oviposition preference was not influenced by differences in leaf size or adherence to leaves by ovipositing moths. The high specific leaf weights common to adult leaves were associated with reduced oviposition.
• 3 Although neonates fed on both juvenile and adult leaves of most of the trees studied, performance was greater on juvenile as opposed to adult foliage. Juvenile leaves typically had lower specific leaf weights and were nutritionally superior to their adult counterparts. Specific leaf weights above 0.236 mg/mm² (associated with low water, i.e. = 56.5%, and nitrogen, i.e. = 1.23%, contents) were associated with reduced larval performance. Younger adult leaves, those with lower specific leaf weights, allowed slightly greater larval consumption.
• 4 When ovipositing, this eucalypt‐specific moth discriminates between leaf types of its heterophyllous hosts in favour of types with the lowest specific leaf weight. Less tough leaf types, which are also higher in nitrogen, enable neonates to attain larger body weights.

     

Geographical variation and population structure in the White-rumped Sandpiper Calidris fuscicollis as shown by morphology, mitochondrial DNA and carbon isotope ratios

Several plant-herbivore hypotheses are based on the assumption that plants cannot simultaneously allocate resources to growth and defence. We studied seasonal patterns in allocation to growth and putatively defensive compounds by monitoring several chemical and physical traits in the leaves of mountain birch from early June (budburst) to late September (leaf senescence). We found significant seasonal changes in all measured characteristics, both in terms of concentrations (mg g^-1) and amounts (mg leaf^-1). Changes were very rapid in the spring, slow in the middle of the season, and there was another period of fast changes in the senescing leaves. Co-occurring changes in physical leaf traits and concentrations of several compounds indicated a seasonal decline in foliage suitability for herbivores. Concentrations of protein and free amino acids declined through the growing season whereas individual sugars showed variable seasonal patterns. The seasonal trends of phenolic groups differed drastically: concentrations of soluble proanthocyanidins increased through the season, whereas cell wall-bound proanthocyanidins, gallotannins and flavonoid glycosides declined after an initial increase in young leaves. We failed to find proof that the seasonal accumulation of phenolics would have been seriously compromised by leaf or shoot growth, as assumed by the growth/differentiation balance hypothesis and the protein competition model hypothesis. On the contrary, there was a steady increase in the total amount of phenolics per leaf even during the most active leaf growth.     

Putting the insect into the birch–insect interaction

Leaf maturation in mountain birch (Betula pubescens ssp. czerepanovii) is characterized by rapid shifts in the types of dominant phenolics: from carbon-economic flavonoids aglycons in flushing leaves, via hydrolysable tannins and flavonoid glycosides, to carbon-rich proanthocyanidins (condensed tannins) in mature foliage. This shift accords with the suggested trade-offs between carbon allocation to plant defense and growth, but may also relate to the simultaneous decline in nutritive leaf traits, such as water, proteins and sugars, which potentially limit insect growth. To elucidate how birch leaf quality translates into insect growth, I introduce a simple model that takes into account defensive compounds but also acknowledges insect demand for nutritive compounds. The effects of defensive compounds on insect growth depend strongly on background variation in nutritive leaf traits: compensatory feeding on low nutritive diets increases the intake of defensive compounds, and the availability of growth-limiting nutritive compounds may modify the effects of defenses. The ratio of consumption to larval growth (both in dry mass) increases very rapidly with leaf maturation: from 2.9 to 9.8 over 2 weeks in June-July, and to 15 by August. High concentrations in mature birch leaves of "quantitative" defenses, such as proanthocyanidins (15-20% of dry mass), presumably prevent further consumption. If the same compounds had also protected half-grown leaves (which supported the same larval growth with only one third of the dry matter consumption of older leaves), the same intake of proanthocyanidins would have demanded improbably high concentrations (close to 50%) in young leaves. The model thus suggests an adaptive explanation for the high levels of "quantitative" defenses, such as proanthocyanidins, in low-nutritive but not in high-nutritive leaves because of the behavioral responses of insect feeding to leaf nutritive levels.     

Through-flow of water in leaves of a submerged plant is influenced by the apical opening

Submerged aquatic higher plants maintain acropetal water transport to the young leaves in active growth to satisfy their demand for nutrients and hormones derived from the roots. We here present the first measurements of hydraulic properties for a submerged plant, the monocotyledon Sparganium emersum Rehman. The hydraulic conductance per unit length, K_h, was measured in leaf segments without the leaf tip and shown to be greater in old, fully developed leaves (1.5 · 10⁻¹⁰ · m⁴ · MPa⁻¹ · s⁻¹) than in young leaves (1.0 · 10⁻¹⁰ · m⁴ · MPa⁻¹ · s⁻¹). In leaves with intact leaf tips, however, K_h was significantly greater in the youngest leaves, which suggests that the leaf tip with the hydathode influences resistance and thus flow. Microscopy confirmed that the hydathodal area, which is an apical opening, undergoes structural changes with leaf age; a matrix of microorganisms develops in the older leaves and probably restricts water flow by clogging the hydathodes. The leaf specific conductivity expressing transport capacity relative to the leaf area supplied, of S. emersum (0.1 · 10⁻⁸ to 9 · 10⁻⁸ · m² MPa⁻¹· s⁻¹) was within the same range as for various species of terrestrial ferns, vines and trees. This finding does not support the traditional concept of functionally reduced vascular transport in Received: 15 July 1996 / Accepted: 30 November 1996     

Dieback of rural eucalypts: Does insect herbivory relate to dietary quality of tree foliage?

Foliar dietary quality, and the damage that insects caused to the foliage of dieback-affected and healthy Eucalyptus blakelyi trees, were monitored for 3 years, on pastoral properties in the Australian Capital Territory. Compared with healthy trees, the foliage of dieback trees was more heavily grazed by insects, and its dietary quality was generally superior. Some of the differences in dietary quality were related to the average age of the foliage of healthy and dieback trees. But when statistical models were used to equalize the effects of differences in leaf ages, leaves on dieback trees nonetheless tended to have lower specific weights, and were sometimes rounder and contained more nitrogen. Regression analyses of herbivory against each of the dietary quality variables showed that the only significant relationship that was consistent for both of the years monitored was for foliar nitrogen and herbivory for both dieback and healthy trees. In the first year, regressions between herbivory and specific leaf weight, shape, or sugar content were also significant, but only amongst the dieback trees. This may indicate that these relationships were a response to, rather than a primary cause of, the repeated high defoliation of the dieback trees. Multiple regression equations incorporating annual means of several quality variables explained a high proportion of the variance in annual herbivory, but were grossly different between years.     

Performance of moth larvae on birch in relation to altitude, climate, host quality and parasitoids

We studied topographical and year-to-year variation in the performance (pupal weights, survival) and larval parasitism of Epirrita autumnata larvae feeding on mountain birch in northernmost Finland in 1993–1996. We found differences in both food plant quality and parasitism between sites ranging from 80 m to 320 m above sea level. Variation in food plant quality had particularly marked effects on larval survival. The advanced phenology of the birches in relation to the start of the larval period reduced pupal weights. Parasitism rates were different between years and between sites. The clearest site differences were in the proportions of different parasitoid species: Eulophus larvarum was most abundant at the lowest-altitude sites, and Cotesia jucunda at the highest. Differences in the performance of E. autumnata were related to temperature conditions: at higher temperatures, survival and the egg production index were lower, and larval parasitism was higher than at lower temperatures. The higher parasitism at higher temperatures was probably due to greater parasitoid activity during warmer days. In the comparison of different sources of spatial and annual variation in the performance of E. autumnata, the most important factor appeared to be egg mortality related to minimum winter temperature, followed by parasitism and, finally, the variation in food plant quality. If, as predicted, the climate gradually warms up, the effects of warmer summers on the outbreaks of E. autumnata suggest a decrease in outbreak intensity. Received: 4 January 1999 / Accepted: 22 March 1999     

Combined effects of elevated CO2 and soil drought on carbon and nitrogen allocation of the desert shrub Caragana intermedia

Impacts of either elevated CO₂ or drought stress on plant growth have been studied extensively, but interactive effects of these on plant carbon and nitrogen allocation is inadequately understood yet. In this study the response of the dominant desert shrub, Caragana intermedia Kuanget H.c.Fu, to the interaction of elevated CO₂ (700 ± 20 μmol mol⁻¹) and soil drought were determined in two large environmental growth chambers (18 m²). Elevated CO₂ increased the allocation of biomass and carbon into roots and the ratio of carbon to nitrogen (C:N) as well as the leaf soluble sugar content, but decreased the allocation of biomass and carbon into leaves, leaf nitrogen and leaf soluble protein concentrations. Elevated CO₂ significantly decreased the partitioning of nitrogen into leaves, but increased that into roots, especially under soil drought. Elevated CO₂ significantly decreased the carbon isotope discrimination (Δ) in leaves, but increased them in roots, and the ratio of Δ values between root and leaf, indicating an increased allocation into below-ground parts. It is concluded that stimulation of plant growth by CO₂ enrichment may be negated under soil drought, and under the future environment, elevated CO₂ may partially offset the negative effects of enhanced drought by regulating the partitioning of carbon and nitrogen.     

Phenolic and phenolic-related factors as determinants of suitability of mountain birch leaves to an herbivorous insect

We investigated the role of phenolic and phenolic-related traits of the leaves of mountain birch (Betula pubescens ssp. czerepanovii) as determinants of their suitability for the growth of larvae of the geometrid Epirrita autumnata. As parameters of leaf suitability, we determined the contents of total phenolics, gallotannins, soluble and cell-wall-bound proanthocyanidins (PAS and PAB, respectively), lignin, protein precipitation capacity of tannins (PPC), and leaf toughness. In addition, we examined concentrations of soluble carbohydrates and protein-bound amino acids as background variables describing the nutritive value of leaves. The correlation of the leaf traits of our 40 study trees with the tree-specific relative growth rate (RGR) of E. autumnata showed that the only significant correlation with RGR was that of PAS - the largest fraction of total phenolics - and even that explained only 15% of the variation in E. autumnata growth. The nonlinear estimation of the relationship between RGR and PAS by piecewise linear regression divided the 40 study trees into two groups: (i) 19 trees with good leaves for E. autumnata (RGR ranging from 0.301 to 0.390), and (ii) 21 trees with poor leaves (RGR ranging from 0.196 to 0.296). The suitability of leaves within these two groups of trees was determined by different phenolic traits. Within the good group, the suitability of leaves for larvae was determined by the PPC of extracts, which strongly correlated with gallotannins, and by the total content of gallotannins. In contrast, the leaves of poor trees had significantly higher contents of both PAS and PAB, but leaf toughness correlated only negatively with the RGR of E. autumnata larvae. We also discuss the causes of variation in the phenolic and phenolic-related factors that determine the suitability of leaves for E. autumnata larvae in different groups of trees.     

Opposing effects of elevated CO2 and N deposition on Lymantria monacha larvae feeding on spruce trees

The effects of elevated atmospheric CO₂ and increased wet N deposition on leaf quality and insect herbivory were evaluated in nine model ecosystems composed of 7-year-old spruce trees (Picea abies) and three understorey species established on natural forest soil. Each model ecosystem was grown in a simulated montane climate, and was exposed to one of three CO₂ concentrations (280, 420, and 560 μl l⁻¹), and to one of three levels of N deposition (0, 30, and 90 kg ha⁻¹ year⁻¹) for 3 years. In the 3rd year of the experiment second to third instars of the nun moth (Lymantria monacha) were allowed to feed directly on current-year needles of top canopy branches of each tree for 12 days. Specific leaf area (SLA), water content, and N concentration decreased in needles exposed to elevated CO₂, whereas the concentrations of starch, condensed tannins, and total phenolics increased. Increased N deposition had no significant effect on SLA, and water content, but the concentrations of starch, condensed tannins, and total phenolics decreased, and sugar and N concentrations increased. Despite higher relative consumption rates (RCRs) larvae consumed 33% less N per unit larval biomass and per day at the two high CO₂ treatments, compared to those feeding on 280 μl l⁻¹-needles, but they maintained similar N accumulation rates due to increased N utilization efficiencies (NUE). However, over the 12-day experimental period larvae gained less N overall and reached a 35% lower biomass in the two high-CO₂ treatments compared to those at 280 μl l⁻¹. The effects of increased N deposition on needle quality and insect performance were generally opposite to those of CO₂ enrichment, but were lower in magnitude. We conclude that altered needle quality in response to elevated CO₂ will impair the growth and development of L. monacha larvae. Increasing N deposition may mitigate these effects, which could lead to altered insect herbivore distributions depending on regional patterns of N deposition. Received: 8 June 1998 / Accepted: 27 October 1998     

The fraction of expanding to expanded leaves determines the biomass response of Populus to elevated CO2

We examined whether the effects of elevated CO₂ on growth of 1-year old Populus deltoides saplings was a function of the assimilation responses of particular leaf developmental stages. Saplings were grown for 100 days at ambient (approximately 350 ppm) and elevated (ambient + 200 ppm) CO₂ in forced-air greenhouses. Biomass, biomass distribution, growth rates, and leaf initiation and expansion rates were unaffected by elevated CO₂. Leaf nitrogen (N), the leaf C:N ratio, and leaf lignin concentrations were also unaffected. Carbon gain was significantly greater in expanding leaves of saplings grown at elevated compared to ambient CO₂. The Rubisco content in expanding leaves was not affected by CO₂ concentration. Carbon gain and Rubisco content were significantly lower in fully expanded leaves of saplings grown at elevated compared to ambient CO₂, indicating CO₂-induced down-regulation in fully expanded leaves. Elevated CO₂ likely had no overall effect on biomass accumulation due to the more rapid decline in carbon gain as leaves matured in saplings grown at elevated compared to ambient CO₂. This decline in carbon gain has been documented in other species and shown to be related to a balance between sink/source balance and acclimation. Our data suggest that variation in growth responses to elevated CO₂ can result from differences in leaf assimilation responses in expanding versus expanded leaves as they develop under elevated CO₂. Received: 28 September 1998 / Accepted: 23 June 1999     

The effects of enriched CO2 atmospheres on plant-insect herbivore interactions: growth responses of larvae of the specialist butterfly,Junonia coenia (Lepidoptera: Nymphalidae)

Summary Little is known about the effects of enriched CO₂ environments, which are anticipated to exist in the next century, on natural plant-insect herbivore interactions. To begin to understand such effects on insect growth and survival, I reared both early and penultimate instar larvae of the buckeye, Junonia coenia (Lepidoptera: Nymphalidae), on leaves from one of their major hostplants, plantain, Plantago lanceolata (Plantaginaceae), grown in either ambient (350 PPM) or high (700 PPM) CO₂ atmospheres. Despite consuming more foliage, early instar larvae experienced reduced growth on high CO₂-grown compared to ambient CO₂-grown leaves. However, survivorship of early instar larvae was unaffected by the CO₂ treatment. Larval weight gain was positively correlated with the nitrogen concentration of the plant material and consumption was negatively correlated with foliar nitrogen concentration, whereas neither larval weight gain nor consumption were significantly correlated with foliar water or allelochemical concentrations. In contrast, penultimate instar larvae had similar growth rates on ambient and high CO₂-grown leaves. Significantly higher consumption rates on high CO₂-grown plants enabled penultimate instar larvae to obtain similar amounts of nitrogen in both treatments. These larvae grew at similar rates on foliage from the two CO₂ treatments, despite a reduced efficiency of conversion of ingested food (ECI) on the low nitrogen, high CO₂-grown plants. However, nitrogen utilization efficiencies (NUE) were unaffected by CO₂ treatment. Again, for late instar larvae, consumption rates were negatively correlated with foliar nitrogen concentrations, and ECI was also very highly correlated with leaf nitrogen; foliar water or allelochemical concentrations did not affect either of these parameters. Differences in growth responses of early and late instar larvae to lower nitrogen, high-CO₂ grown foliage may be due to the inability of early instar larvae to efficiently process the increased flow of food through the gut caused by additional consumption of high CO₂ foliage.