Perfect is best: low leaf fluctuating asymmetry reduces herbivory by leaf miners

Fluctuating asymmetry (FA) represents small, random variation from symmetry and can be used as an indicator of plant susceptibility to herbivory. We investigated the effects of FA of two oak species, Quercus laevis and Q. geminata, and the responses of three herbivore guilds: leaf miners, gallers, and chewers. To examine differences in FA and herbivory between individuals, 40 leaves from each tree were collected, and FA indices were calculated. To examine differences in FA and herbivory within-individuals, we sampled pairs of mined and unmined leaves for asymmetry measurements. Differences in growth of leaf miners between leaf types were determined by tracing 50 mines of each species on symmetric leaves and asymmetric leaves. Asymmetric leaves contained significantly lower concentrations of tannins and higher concentrations of nitrogen than symmetric leaves for both plant species. Both frequency of asymmetric leaves on plants and levels of asymmetry positively influenced the abundance of Brachys, Stilbosis and other leaf miners, but no significant relationship between asymmetry and herbivory was observed for Acrocercops. Brachys and Stilbosis mines were smaller on asymmetric leaves, but differences in mine survivorship between symmetric and asymmetric leaves were observed only for Stilbosis mines. This study indicated that leaf miners might use leaf FA as a cue to plant quality, although differential survivorship among leaf types was not observed for all species studied. Reasons for the different results between guilds are discussed.     

Seeing the forest for the trees: long-term exposure to elevated CO2 increases some herbivore densities

The effects of elevated CO₂ on plant growth and insect herbivory have been frequently investigated over the past 20 years. Most studies have shown an increase in plant growth, a decrease in plant nitrogen concentration, an increase in plant secondary metabolites and a decrease in herbivory. However, such studies have generally overlooked the fact that increases in plant production could cause increases of herbivores per unit area of habitat. Our study investigated leaf production, herbivory levels and herbivore abundance per unit area of leaf litter in a scrub‐oak system at Kennedy Space Center, Florida, under conditions of ambient and elevated CO₂, over an 11‐year period, from 1996 to 2007. In every year, herbivory, that is leafminer and leaftier abundance per 200 leaves, was lower under elevated CO₂ than ambient CO₂ for each of three species of oaks, Quercus myrtifolia, Quercus chapmanii and Quercus geminata. However, leaf litter production per 0.1143 m² was greater under elevated CO₂ than ambient CO₂ for Q. myrtifolia and Q. chapmanii, and this difference increased over the 11 years of the study. Leaf production of Q. geminata under elevated CO₂ did not increase. Leafminer densities per 0.1143 m² of litterfall for Q. myrtifolia and Q. chapmanii were initially lower under elevated CO₂. However, shortly after canopy closure in 2001, leafminer densities per 0.1143 m² of litter fall became higher under elevated CO₂ and remained higher for the remainder of the experiment. Leaftier densities per 0.1143 m² were also higher under elevated CO₂ for Q. myrtifolia and Q. chapmanii over the last 6 years of the experiment. There were no differences in leafminer or leaftier densities per 0.1143 m² of litter for Q. geminata. These results show three phenomena. First, they show that elevated CO₂ decreases herbivory on all oak species in the Florida scrub‐oak system. Second, despite lower numbers of herbivores per 200 leaves in elevated CO₂, increased leaf production resulted in higher herbivore densities per unit area of leaf litter for two oak species. Third, they corroborate other studies which suggest that the effects of elevated CO₂ on herbivores are species specific, meaning they depend on the particular plant species involved. Two oak species showed increases in leaf production and herbivore densities per 0.1143 m² in elevated CO₂ over time while another oak species did not. Our results point to a future world of elevated CO₂ where, despite lower plant herbivory, some insect herbivores may become more common.     

Acclimation of photosynthesis and respiration to elevated atmospheric CO2 in two Scrub Oaks

Abstract For two species of oak, we determined whether increasing atmospheric CO₂ concentration (C_a) would decrease leaf mitochondrial respiration (R) directly, or indirectly owing to their growth in elevated C_a, or both. In particular, we tested whether acclimatory decreases in leaf‐Rubisco content in elevated C_a would decrease R associated with its maintenance. This hypothesis was tested in summer 2000 on sun and shade leaves of Quercus myrtifolia Willd. and Quercus geminata Small. We also measured R on five occasions between summer 1999 and 2000 on leaves of Q. myrtifolia. The oaks were grown in the field for 4 years, in either current ambient or elevated (current ambient + 350 µmol mol^?1) C_a, in open‐top chambers (OTCs). For Q. myrtifolia, an increase in C_a from 360 to 710 µmol mol^?1 had no direct effect on R at any time during the year. In April 1999, R in young Q. myrtifolia leaves was significantly higher in elevated C_a—the only evidence for an indirect effect of growth in elevated C_a. Leaf R was significantly correlated with leaf nitrogen (N) concentration for the sun and shade leaves of both the species of oak. Acclimation of photosynthesis in elevated C_a significantly reduced maximum RuBP‐saturated carboxylation capacity (V_{c max}) for both the sun and shade leaves of only Q. geminata. However, we estimated that only 11–12% of total leaf N was invested in Rubisco; consequently, acclimation in this plant resulted in a small effect on N and an insignificant effect on R. In this study measurements of respiration and photosynthesis were made on material removed from the field; this procedure had no effect on gas exchange properties. The findings of this study were applicable to R expressed either per unit leaf area or unit dry weight, and did not support the hypothesis that elevated C_a decreases R directly, or indirectly owing to acclimatory decreases in Rubisco content.     

Differential effects of elevated CO2 on acorn density, weight, germination, and predation among three oak species in a scrub-oak forest

Much research on the effects of elevated CO₂ on forest trees has focused on quantitative changes in photosynthesis, secondary chemistry, and plant biomass. However, plant fitness responses to rising CO₂ should also include quantitative measures of reproduction, since most forest systems are recruitment limited. Until now, it has proved very difficult to grow forest trees to sexual maturity in a CO₂‐enriched environment. This paper is the first of its kind to address the effects of elevated CO₂ on the reproduction of hardwood trees in a natural forest. Beginning in 1996, scrub‐oak vegetation, predominantly three species of scrub‐oaks, Quercus myrtifolia, Q. chapmanii, and Q. geminata, were grown inside eight chambers with elevated CO₂ (704 parts per million (ppm)) and eight with ambient CO₂ (379 ppm) at Kennedy Space Center, Florida. In elevated CO₂, acorn production increased significantly for the dominant species Q. myrtifolia and for Q. chapmanii, but it did not increase for the subdominant, Q. geminata. Acorn weight, germination rate, and predation by weevils were unaffected by CO₂. Thus, recruitment of some forest tree species into the Florida scrub‐oak community is likely to be accelerated in an atmosphere of increased CO₂. However, because the acceleration of recruitment differs among species, over the long term, Q. myrtifolia and Q. chapmanii will be favored over Q. geminata and this is likely to change patterns of species diversity.     

Similar responses of insect herbivores to leaf fluctuating asymmetry

Fluctuating asymmetry (FA) represents small, random variation from symmetry and it has been used as an indicator of plant quality and susceptibility to herbivory. In this study, the effects of FA on the responses of distinct herbivore species belonging to several guilds were examined along an environmental gradient in south Florida. This approach was chosen because it relies on a multi-species approach to the study of fluctuating asymmetry and patterns of herbivory between and within plants along an environmental gradient of salinity and plant stress. To examine differences in FA between and within plant communities, seven plant species were investigated. Four of these plants were coastal species and three species occurred in upland communities. Levels of FA were assessed before herbivory and plants were followed for the whole herbivory season in 2006. Coastal plants exhibited significantly higher salt concentration, higher percentage of asymmetric leaves and higher asymmetry levels than upland plants. Herbivore abundance varied widely amongst the seven species studied, but quantitative syntheses of our results indicated significant and positive responses of insect herbivores to leaf asymmetry: insects were 25.11% more abundant on more asymmetric plants and stronger effects of asymmetry were observed for leaf miners compared to gall-formers. As demonstrated by other recent studies, FA might be used as a reliable stress indicator, leading to similar responses of insect herbivores to variation in leaf symmetry.     

Elevated atmospheric CO2 stimulates aboveground biomass in a fire-regenerated scrub-oak ecosystem

The effect of elevated atmospheric CO₂ concentration (C_a) on the aboveground biomass of three oak species, Quercus myrtifolia, Q. geminata, and Q. chapmanii, was estimated nondestructively using allometric relationships between stem diameter and aboveground biomass after four years of experimental treatment in a naturally fire‐regenerated scrub‐oak ecosystem. After burning a stand of scrub‐oak vegetation, re‐growing plants were exposed to either current ambient (379 µL L^?1 CO₂) or elevated (704 µL L^?1 CO₂) C_a in 16 open‐top chambers over a four‐year period, and measurements of stem diameter were carried out annually on all oak shoots within each chamber. Elevated C_a significantly increased aboveground biomass, expressed either per unit ground area or per shoot; elevated C_a had no effect on shoot density. The relative effect of elevated C_a on aboveground biomass increased each year of the study from 44% (May 96–Jan 97), to 55% (Jan 97–Jan 98), 66% (Jan 98–Jan 99), and 75% (Jan 99–Jan 00). The effect of elevated C_a was species specific: elevated C_a significantly increased aboveground biomass of the dominant species, Q. myrtifolia, and tended to increase aboveground biomass of Q. chapmanii, but had no effect on aboveground biomass of the subdominant, Q. geminata. These results show that rising atmospheric CO₂ has the potential to stimulate aboveground biomass production in ecosystems dominated by woody species, and that species‐specific growth responses could, in the long term, alter the composition of the scrub‐oak community.     

Plastic leaf morphology in three species of Quercus: The more exposed leaves are smaller,more lobated and denser

Phenotypic plasticity and developmental instability in leaf traits are common in oak species but the role of environmental factors is not well understood. To decipher possible correlations between different leaf traits and effects of the position of leaves within the tree canopy, we quantified the plasticity of three leaf traits of 30 trees of Quercus alba L., Quercus palustris Muench and Quercus velutina Lam. We hypothesized that trees could modify the shape of their leaves for better adaptation to the variable microclimate within the canopy. Our results demonstrated that the south and north outer leaves were significantly smaller, more lobed and denser than those situated in the inner canopy. The order of leaves on the branch accounted for the plasticity of leaf traits in Q. alba only. Plasticity of lobing in Q. alba and Q. velutina depended on the height of the trees. We detected fluctuating asymmetry (FA) in all three species, but the source of variation depended on branch position in Q. velutina only. FA was more pronounced in north-facing leaves. Plasticity of the leaf traits ranged from small to medium. Plasticity of leaf area and leaf mass per area (LMA) depended on the branch position. However, the plasticity of lobation was not affected by the location of a branch within the tree canopy. Quercus alba and Q. palustris had similar plastic responses but the plasticity of Q. velutina was significantly smaller. We concluded that individual plants detect and cope with environmental stress through vegetative organ modification.     

Differential resource utilization and co-occurrence of leaf miners on oak (Quercus dentata)

Abstract.

• 1 Spatial, temporal, and dietary differences in resource utilization and patterns of interspecific association on leaves were investigated for dominant and common leaf-mining species on an oak species, Quercus dentata Thunb., in Hokkaido, northern Japan.
• 2 Leaf miners were divided into two groups according to leaf tissues used for food: upper-layer-feeders which consume the palisade paren-chymatous layer (Stigrnella spp. and tenthredinid sp.), and full-depth-feeders which consume spongy and palisade parenchymatous layers (Phyllonorycter leucocorona (Kumata), P.similis Kumata, and Caloptilia sapporella (Matsumura)).
• 3 Differences in the position of mines on leaves were found among species: mines of P.similis were distributed more frequently in the middle section of leaves, whereas those of the remaining species were concentrated in the basal section.
• 4 Leaf size preference differed between species: C.sapporella and tenthredinid sp. tended to select larger leaves more frequently than did the other species.
• 5 Phenological differentiation was found among species: C.sapporella appeared earliest, followed by P.similis, P.leucocorona and a tenthredinid sp., and then Stigmella spp.
• 6 Each species showed a highly clumped distribution among leaves. Leaf miners of some species pairs co-occurred on leaves more frequently than expected by chance.

     

Impacts of urbanization on insect herbivory and plant defences in oak trees

Systematic comparisons of species interactions in urban versus rural environments can improve our understanding of shifts in ecological processes due to urbanization. However, such studies are relatively uncommon and the mechanisms driving urbanization effects on species interactions (e.g. between plants and insect herbivores) remain elusive. Here we investigated the effects of urbanization on leaf herbivory by insect chewers and miners associated with the English oak Quercus robur by sampling trees in rural and urban areas throughout most of the latitudinal distribution of this species. In performing these comparisons, we also controlled for the size of the urban areas (18 cities) and gathered data on CO₂ emissions. In addition, we assessed whether urbanization affected leaf chemical defences (phenolic compounds) and nutritional traits (phosphorus and nitrogen), and whether such changes correlated with herbivory levels. Urbanization significantly reduced leaf chewer damage but did not affect leaf miners. In addition, we found that leaves from urban locations had lower levels of chemical defences (condensed and hydrolysable tannins) and higher levels of nutrients (nitrogen and phosphorus) compared to leaves in rural locations. The magnitude of urbanization effects on herbivory and leaf defences was not contingent upon city size. Importantly, while the effects of urbanization on chemical defences were associated with CO₂ emissions, changes in leaf chewer damage were not associated with either leaf traits or CO₂ levels. These results suggest that effects of urbanization on herbivory occur through mechanisms other than changes in the plant traits measured here. Overall, our simultaneous assessment of insect herbivory, plant traits and abiotic correlates advances our understanding of the main drivers of urbanization effects on plant–herbivore interactions.     

The role of leaf and canopy-level gas exchange in the replacement of Quercus virginiana (Fagaceae) by Juniperus ashei (Cupressaceae) in semiarid savannas

Photosynthesis, transpiration, and leaf area distribution were sampled in mature Quercus virginiana and Juniperus ashei trees to determine the impact of leaf position on canopy-level gas exchange, and how gas exchange patterns may affect the successful invasion of Quercus communities by J. ashei. Sampling was conducted monthly over a 2-yr period in 12 canopy locations (three canopy layers and four cardinal directions). Photosynthetic and transpiration rates of both species were greatest in the upper canopy and decreased with canopy depth. Leaf photosynthetic and transpiration rates were significantly higher for Q. virginiana (4.1–6.7 μmol CO₂·m⁻²·s⁻¹ and 1.1–2.1 mmol H₂O·m⁻²·s⁻¹) than for J. ashei (2.1–2.8 μmol CO₂·m⁻²·s⁻¹ and 0.7–1.0 mmol H₂O·m⁻²·s⁻¹) in every canopy level and direction. Leaves on the south and east sides of both species had higher gas exchange rates than leaves on the north and west sides. Although Quercus had a greater mean canopy diameter than Juniperus (31.3 vs. 27.7 m²), J. ashei had significantly greater leaf area (142 vs. 58 m²/tree). A simple model combining leaf area and gas exchange rates for different leaf positions demonstrated a significantly greater total canopy carbon dioxide uptake for J. ashei compared to Q. virginiana (831 vs. 612 g CO₂·tree⁻¹·d⁻¹, respectively). Total daily water loss was also greater for Juniperus (125 vs. 73 Ltree⁻¹·d⁻¹). Differences in leaf gas exchange rates were poor predictors of the relationship between the invasive J. ashei and the codominant Q. virginiana. Leaf area and leaf area distribution coupled with leaf gas exchange rates were necessary to demonstrate the higher overall competitive potential of J. ashei.     

Consequences of CO2 and light interactions for leaf phenology, growth, and senescence in Quercus rubra

We investigated how light and CO₂ levels interact to influence growth, phenology, and the physiological processes involved in leaf senescence in red oak (Quercus rubra) seedlings. We grew plants in high and low light and in elevated and ambient CO₂. At the end of three years of growth, shade plants showed greater biomass enhancement under elevated CO₂ than sun plants. We attribute this difference to an increase in leaf area ratio (LAR) in shade plants relative to sun plants, as well as to an ontogenetic effect: as plants increased in size, the LAR declined concomitant with a decline in biomass enhancement under elevated CO₂ Elevated CO₂ prolonged the carbon gain capacity of shade‐grown plants during autumnal senescence, thus increasing their functional leaf lifespan. The prolongation of carbon assimilation, however, did not account for the increased growth enhancement in shade plants under elevated CO₂. Elevated CO₂ did not significantly alter leaf phenology. Nitrogen concentrations in both green and senesced leaves were lower under elevated CO₂ and declined more rapidly in sun leaves than in shade leaves. Similar to nitrogen concentration, the initial slope of A/C_i curves indicated that Rubisco activity declined more rapidly in sun plants than in shade plants, particularly under elevated CO₂. Absolute levels of chlorophyll were affected by the interaction of CO₂ and light, and chlorophyll content declined to a minimal level in sun plants sooner than in shade plants. These declines in N concentration, in the initial slope of A/C_i curves, and in chlorophyll content were consistent with declining photosynthesis, such that elevated CO₂ accelerated senescence in sun plants and prolonged leaf function in shade plants. These results have implications for the carbon economy of seedlings and the regeneration of red oak under global change conditions.     

Leaf trait variation on Erythroxylum tortuosum (Erythroxylaceae) and its relationship with oviposition preference and stress by a host‐specific leaf miner

It has been suggested that plant physical and chemical traits vary considerably in space and time. Hence, leaf‐mining insects may adjust their oviposition in response to leaf attributes representing high quality. Moreover, herbivorous insects can modify leaf morphology by acting as stressors, increasing, for example, fluctuating asymmetry (FA) levels. Here, we investigate oviposition preference in Agnippe sp.2, a leaf‐mining moth of Erythroxylum tortuosum, in relation to differences in leaf nutritional quality (i.e. levels of water, nitrogen and tannin content), leaf area (i.e. quantity of resource hypothesis) and FA. We also verify whether temporal variation in plant nutritional quality emerges as an alternative hypothesis to explain oviposition distribution in time, and whether this leaf miner is a stress‐causing agent, increasing FA during larval development. Mined leaves and leaves with and without eggs were periodically collected from plants located in a Cerrado fragment in Brazil. In the laboratory, leaf traits were assessed (using image analysis software) and quantified (biochemical analysis) according to the aims previously determined. Oviposition probability did not change in relation to variations in nitrogen, tannins and FA of leaves. However, leaf‐miner females preferred to oviposit on leaves having large areas and low water contents. It was also verified that new leaves of E. tortuosum, which carried most leaf‐miner eggs, presented significantly lower tannins and greater levels of nitrogen and water than old leaves. The oviposition choice exhibited by leaf miners was found to be non‐random because they appear to use resource quantity and water content as cues as where to lay their eggs. The temporal variation of plant nutritional quality is likely to influence the time of leaf‐miner oviposition; and leaf FA was not increased during larval feeding, suggesting that these herbivores do not cause variations in FA levels.     

Leaf senescence of Quercus myrtifolia as affected by long-term CO2 enrichment in its native environment

The long‐term effects of elevated (ambient plus 350 μmol mol^?1) atmospheric CO₂ concentration (C_a) on the leaf senescence of Quercus myrtifolia Willd was studied in a scrub‐oak community during the transition from autumn (December 1997) to spring (April 1998). Plants were grown in large open‐top chambers at the Smithsonian CO₂ Research Site, Merritt Island Wildlife Refuge, Cape Canaveral, Florida. Chlorophyll (a + b) concentration, Rubisco activity and N concentration decreased by 75%, 82%, and 52%, respectively, from December (1997) to April (1998) in the leaves grown at ambient C_a. In contrast, the leaves of plants grown at elevated C_a showed no significant decrease in chlorophyll (a + b) concentration or Rubisco activity, and only a 25% reduction in nitrogen. These results indicate that leaf senescence was delayed during this period at elevated C_a. Delayed leaf senescence in elevated C_a had important consequences for leaf photosynthesis. In elevated C_a the net photosynthetic rate of leaves that flushed in Spring 1997 (last year's leaves) and were 13 months old was not different from fully‐expanded leaves that flushed in 1998, and were approximately 1 month old (current year's leaves). In ambient C_a the net photosynthetic rate of last year's leaves was 54% lower than for current year's leaves. When leaves were fully senesced, nitrogen concentration decreased to about 40% of the concentration in non‐senesced leaves, in both CO₂ treatments. In April, net photosynthesis was 97% greater in leaves grown in elevated C_a than in those grown at ambient. During the period when elevated C_a delayed leaf senescence, more leaves operating at higher photosynthetic rate would allow the ecosystem dominated by Q. myrtifolia to gain more carbon at elevated C_a than at ambient C_a.     

Elevated CO2 increases the long-term decomposition rate of Quercus myrtifolia leaf litter

Decomposition of Quercus myrtifolia leaf litter in a Florida scrub oak community was followed for 3 years in two separate experiments. In the first experiment, we examined the effects CO₂ and herbivore damage on litter quality and subsequent decomposition. Undamaged, chewed and mined litter generated under ambient and elevated (ambient+350 ppm V) CO₂ was allowed to decompose under ambient conditions for 3 years. Initial litter chemistry indicated that CO₂ levels had minor effects on litter quality. Litter damaged by leaf miners had higher initial concentrations of condensed tannins and nitrogen (N) and lower concentrations of hemicellulose and C : N ratios compared with undamaged and chewed litter. Despite variation in litter quality associated with CO₂, herbivory, and their interaction, there was no subsequent effect on rates of decomposition under ambient atmospheric conditions. In the second experiment, we examined the effects of source (ambient and elevated) of litter and decomposition site (ambient and elevated) on litter decomposition and N dynamics. Litter was not separated by damage type. The litter from both elevated and ambient CO₂ was then decomposed in both elevated and ambient CO₂ chambers. Initial litter chemistry indicated that concentrations of carbon (C), hemicellulose, and lignin were higher in litter from elevated than ambient CO₂ chambers. Despite differences in C and fiber concentrations, litter from ambient and elevated CO₂ decomposed at comparable rates. However, the atmosphere in which the decomposition took place resulted in significant differences in rates of decomposition. Litter decomposing under elevated CO₂ decomposed more rapidly than litter under ambient CO₂, and exhibited higher rates of mineral N accumulation. The results suggest that the atmospheric conditions during the decomposition process have a greater impact on rates of decomposition and N cycling than do the atmospheric conditions under which the foliage was produced.     

云丘山不同海拔梯度橿子栎叶性特征研究

在吕梁山南段云丘山,以橿子栎为主要研究对象,采用野外调查和室内实验相结合的方法,对比叶面积(SLA)、叶面积(LA)、叶干物质含量(LDMC)、叶长宽比(L/W)、叶绿素含量(Chl)以及叶氮含量等有代表性的指标进行分析.结果表明：(1)随着海拔梯度的增加,橿子栎 L/W、Chl、SLA、LA、单位面积的叶氮含量(N area )、单位重量的叶氮含量(N mass )、叶饱和鲜重和叶干重均呈先上升后下降的趋势,在海拔1180 m 处有最大值;LDMC 与海拔呈显著负相关(P ＜0．05);(2)橿子栎 LA 与 SLA 和 N mass 呈极显著正相关关系(P ＜0．01),与 LDMC 呈显著负相关关系(P ＜0．05),与 N area呈极显著负相关关系(P ＜0．01);(3)橿子栎叶片 SLA与 L/W 呈显著正相关(P ＜0．05),与 N mass 呈极显著正相关关系(P ＜0．01),橿子栎叶片 SLA 与 LDMC 和N area呈显著负相关关系(P ＜0．05);(4)橿子栎 LDMC 与 N area 呈显著正相关(P ＜0．05).通过对云丘山景区橿子栎叶片性状随海拔变化规律的研究,可以探索橿子栎叶片性状与海拔梯度变化的关系以及最适生长环境,从而为景区橿子栎林的保护和抚育提供理论依据,为合理开发橿子栎林资源提供基础资料.     

Elevated atmospheric CO2 lowers herbivore abundance, but increases leaf abscission rates

Increased levels of atmospheric carbon dioxide (CO₂) are likely to affect the trophic relationships that exist between plants, their herbivores and the herbivores' natural enemies. This study takes advantage of an open‐top CO₂ fertilization experiment in a Florida scrub oak community at Kennedy Space Center, Florida, consisting of eight chambers supplied with ambient CO₂ (360 ppm) and eight chambers supplied with elevated CO₂ (710 ppm). We examined the effects of elevated CO₂ on herbivore densities and levels of leaf consumption, rates of herbivore attack by natural enemies and effects on leaf abscission. Cumulative levels of herbivores and herbivore damage were significantly lower in elevated CO₂ than in ambient CO₂. This may be because leaf nitrogen levels are lower in elevated CO₂. More herbivores die of host plant‐induced death in elevated CO₂ than in ambient CO₂. Attack rates of herbivores by parasitoids are also higher in elevated CO₂, possibly because herbivores need to feed for a longer time in order to accrue sufficient nitrogen (N), thus exposing themselves longer to natural enemies. Insect herbivores cause an increase in abscission rates of leaves throughout the year. Because of the lower insect density in elevated CO₂, we thought, abscission rates would be lower in these chambers. However, abscission rates were significantly higher in elevated CO₂. Thus, the direct effects of elevated CO₂ on abscission are greater than the indirect effects on abscission mediated via lower insect densities. A consequence of increased leaf abscission in elevated CO₂ is that nutrient deposition rates to the soil surface are accelerated.     

Leaf age effects of elevated CO2-grown white oak leaves on spring-feeding lepidopterans

Folivorous insect responses to elevated CO₂-grown tree species may be complicated by phytochemical changes as leaves age. For example, young expanding leaves in tree species may be less affected by enriched CO₂-alterations in leaf phytochemistry than older mature leaves due to shorter exposure times to elevated CO₂ atmospheres. This, in turn, could result in different effects on early vs. late instar larvae of herbivorous insects. To address this, seedlings of white oak (Quercus alba L.), grown in open-top chambers under ambient and elevated CO₂, were fed to two important early spring feeding herbivores; gypsy moth (Lymantria dispar L.), and forest tent caterpillar (Malacosoma disstria Hübner). Young, expanding leaves were presented to early instar larvae, and older fully expanded or mature leaves to late instar larvae. Young leaves had significantly lower leaf nitrogen content and significantly higher total nonstructural carbohydrate:nitrogen ratio as plant CO₂ concentration rose, while nonstructural carbohydrates and total carbon-based phenolics were unaffected by plant CO₂ treatment. These phytochemical changes contributed to a significant reduction in the growth rate of early instar gypsy moth larvae, while growth rates of forest tent caterpillar were unaffected. The differences in insect responses were attributed to an increase in the nitrogen utilization efficiency (NUE) of early instar forest tent caterpillar larvae feeding on elevated CO₂-grown leaves, while early instar gypsy moth larval NUE remained unchanged among the treatments. Later instar larvae of both insect species experienced larger reductions in foliage quality on elevated CO₂-grown leaves than earlier instars, as the carbohydrate:nitrogen ratio of leaves substantially increased. Despite this, neither insect species exhibited changes in growth or consumption rates between CO₂ treatments in the later instar. An increase in NUE was apparently responsible for offsetting reduced foliar nitrogen for the late instar larvae of both species.     

Nutritional quality of leaf detritus altered by elevated atmospheric CO2: effects on development of mosquito larvae

1. Populus tremuloides leaf litter was produced under elevated (ELEV = 720 ppm) and ambient (AMB = 360 ppm) atmospheric CO₂ conditions. Leaf chemical quality was significantly altered by CO₂ enrichment. ELEV leaves had significantly higher concentrations of phenolic compounds and lignins, and higher C : N ratios than AMB. 2. Leaf litter was incubated in a headwater stream for 14 days to become colonised by microorganisms; aquatic bacterial productivity was significantly lower on ELEV than on AMB leaf litter. Colonised leaves were fed to four species of detritivorous mosquito larvae to assess their survivorship and development rates. 3. Larval mortality was 2.2 times higher for Aedes albopictus fed ELEV litter when compared with AMB. Although mortality of A. triseriatus, A. aegypti and Armigeres subalbatus was not affected by treatment, larval development rate was delayed by 78, 25 and 27%, respectively, when fed ELEV litter. 4. Increased mosquito mortality and/or delayed larval development rates are more likely to have negative implications for food web structure and productivity in ecosystems where immature stages of mosquitoes are an important food source of predators.     

Zinc mediated effects on leaf CO2 diffusion conductances and net photosynthesis in Phaseolus vulgaris L.

Supra-optimal levels of zinc in primary leaves of Phaseolus vulgaris increased the CO₂ compensation point and inhibited net photosynthesis. Leaf morphology was modified: mesophyll intercellular area, stomatal slit length and interstomatal distance were reduced, but stomatal density increased. Internal and stomatal conductances to CO₂ diffusion decreased. These changes are discussed in relation to the observed effects on leaf gas exchange and to the previously reported inhibition of different photosynthetic and photorespiratory enzymes.     

Leaf phenology mediates provenance differences in herbivore populations on valley oaks in a common garden

1. Plants from different populations often display a variation in herbivore resistance. However, it is rarely understood what plant traits mediate such differences. 2. It was tested how leaf phenology affects herbivore populations in a 15‐year‐old common garden of valley oaks (Quercus lobata Née) with different populations and maternal parents from throughout the Q. lobata range. 3. The abundance of leaf miners (Stigmella sp. Shrank) and leaf phenology of oaks in the common garden was measured. 4. Leaf miner abundance varied among provenance locations (population), but not among maternal parents within populations. Leaf phenology varied by provenance location and maternal parent, and trees that leafed out earlier accrued higher leaf‐miner abundance. Path analysis indicated that leaf phenology was the likely driver of provenance and parental differences in resistance to leaf miners. 5. Understanding population differences is particularly important when considering transport of genotypes for ornamental or restoration purposes. The present study suggests that similarity in leaf phenology may be one factor that could be used to find genotypes with a similar herbivore resistance to local genotypes.