This review examines the direct effects of climate change on insect herbivores. Temperature is identified as the dominant abiotic factor directly affecting herbivorous insects. There is little evidence of any direct effects of CO2 or UVB. Direct impacts of precipitation have been largely neglected in current research on climate change. Temperature directly affects development, survival, range and abundance. Species with a large geographical range will tend to be less affected. The main effect of temperature in temperate regions is to influence winter survival; at more northerly latitudes, higher temperatures extend the summer season, increasing the available thermal budget for growth and reproduction. Photoperiod is the dominant cue for the seasonal synchrony of temperate insects, but their thermal requirements may differ at different times of year. Interactions between photoperiod and temperature determine phenology; the two factors do not necessarily operate in tandem. Insect herbivores show a number of distinct life‐history strategies to exploit plants with different growth forms and strategies, which will be differentially affected by climate warming. There are still many challenges facing biologists in predicting and monitoring the impacts of climate change. Future research needs to consider insect herbivore phenotypic and genotypic flexibility, their responses to global change parameters operating in concert, and awareness that some patterns may only become apparent in the longer term.

Can heritable traits in a single species affect an entire ecosystem? Recent studies show that such traits in a common tree have predictable effects on community structure and ecosystem processes. Because these 'community and ecosystem phenotypes' have a genetic basis and are heritable, we can begin to apply the principles of population and quantitative genetics to place the study of complex communities and ecosystems within an evolutionary framework. This framework could allow us to understand, for the first time, the genetic basis of ecosystem processes, and the effect of such phenomena as climate change and introduced transgenic organisms on entire communities.

We present evidence that the heritable genetic variation within individual species, especially dominant and keystone species, has community and ecosystem consequences. These consequences represent extended phenotypes, i.e., the effects of genes at levels higher than the population. Using diverse examples from microbes to vertebrates, we demonstrate that the extended phenotype can be traced from the individuals possessing the trait, to the community, and to ecosystem processes such as leaf litter decomposition and N mineralization. In our development of a community genetics perspective, we focus on intraspecific genetic variation because the extended phenotypes of these genes can be passed from one generation to the next, which provides a mechanism for heritability. In support of this view, common‐garden experiments using synthetic crosses of a dominant tree show that their progeny tend to support arthropod communities that resemble those of their parents. We also argue that the combined interactions of extended phenotypes contribute to the among‐community variance in the traits of individuals within communities. The genetic factors underlying this among‐community variance in trait expression, particularly those involving genetic interactions among species, constitute community heritability. These findings have diverse implications. (1) They provide a genetic framework for understanding community structure and ecosystem processes. The effects of extended phenotypes at these higher levels need not be diffuse; they may be direct or may act in relatively few steps, which enhances our ability to detect and predict their effects. (2) From a conservation perspective, we introduce the concept of the minimum viable interacting population (MVIP), which represents the size of a population needed to maintain genetic diversity at levels required by other interacting species in the community. (3) Genotype × environment interactions in dominant and keystone species can shift extended phenotypes to have unexpected consequences at community and ecosystem levels, an issue that is especially important as it relates to global change. (4) Documenting community heritability justifies a community genetics perspective and is an essential first step in demonstrating community evolution. (5) Community genetics requires and promotes an integrative approach, from genes to ecosystems, that is necessary for the marriage of ecology and genetics. Few studies span from genes to ecosystems, but such integration is probably essential for understanding the natural world. Corresponding Editor: A. A. Agrawal

Although rising levels of atmospheric carbon dioxide are expected to directly affect forest ecosystems, little is known of how specific ecological interactions will be modified. This research evaluated the effects of enriched C0 2 on the productivity and phytochemistry of forest trees and performance of associated insects. Our experimental system consisted of three tree species (quaking aspen [Populus tremuloides], red oak [Quercus rubra], sugar maple [Acer saccharum]) that span a range from fast to slow growing, and two species ofleaf-feeding insects (gypsy moth [Lymantria dispar] and forest tent caterpillar [Malacosoma disstria]). Carbon-nutrient balance theory provided a framework for tests of three hypotheses; in response to enriched C0 2 : (I) relative increases in tree growth rates will be greatest for aspen and least for maple, (2) relative decreases in protein and increases in carbon-based compounds will be greatest for aspen and least for maple, and (3) relative reductions in performance will be greatest for insects fed aspen and least for insects fed maple. We grew 1-yr-old seedlings for 60 d under ambient (385 ± 5 J.I.L/ L) or elevated (642 ± 2 J.I.L/L) C0 2 regimes at the University of Wisconsin Biotron. After 50 d, we conducted feeding trials with penultimate-instar gypsy moth and forest tent caterpillars. After 60 d, a second set of trees was harvested and partitioned into root, stem, and leaf tissues. We subsequently analyzed leaf material for a variety of compounds known to affect performance of insect herbivores.In terms of actual dry-matter production, aspen responded the most to enriched C0 2 atmospheres whereas maple responded the least. Proportional growth increases (relative to ambient plants), however, were highest for oak and least for maple. Effects of elevated C0 2 on biomass allocation patterns differed among the three species; root-to-shoot ratios increased in aspen, decreased in oak, and did not change in maple. Enriched C0 2 altered concentrations of primary and secondary metabolites in leaves, but the magnitude and direction of effects were species-specific. Aspen showed the largest change in storage carbon compounds (starch), whereas maple experienced the largest change in defensive carbon compounds (condensed and hydrolyzable tannins). Consumption rates of insects fed high-C0 2 aspen increased dramatically, but growth rates declined. The two species of insects differed in response to oak and maple grown under enriched C0 2 • Gypsy moths grew better on high-C0 2 oak, whereas forest tent caterpillars were unaffected; tent caterpillars tended to grow less on high-C0 2 maple, whereas gypsy moths were unaffected. Changes in insect performance parameters were related to changes in foliar chemistry. Responses of plants and insects agreed with some, but not all, of the predictions of carbon-nutrient balance theory.This study illustrates that tree productivity and chemistry, and the performance of associated insects, will change under C0 2 atmospheres predicted for the next century. Change...

Summary• Salicin-based phenolic glycosides, hydroxycinnamate derivatives and flavonoidderived condensed tannins comprise up to one-third of Populus leaf dry mass. Genes regulating the abundance and chemical diversity of these substances have not been comprehensively analysed in tree species exhibiting this metabolically demanding level of phenolic metabolism.• Here, shikimate-phenylpropanoid pathway genes thought to give rise to these phenolic products were annotated from the Populus genome, their expression assessed by semiquantitative or quantitative reverse transcription polymerase chain reaction (PCR), and metabolic evidence for function presented.• Unlike Arabidopsis , Populus leaves accumulate an array of hydroxycinnamoylquinate esters, which is consistent with broadened function of the expanded hydroxycinnamoyl-CoA transferase gene family. Greater flavonoid pathway diversity is also represented, and flavonoid gene families are larger. Consistent with expanded pathway function, most of these genes were upregulated during wound-stimulated condensed tannin synthesis in leaves.• The suite of Populus genes regulating phenylpropanoid product accumulation should have important application in managing phenolic carbon pools in relation to climate change and global carbon cycling.

Prominent among the many factors now affecting the sustainability of forest ecosystems are anthropogenically-generated carbon dioxide (CO2) and ozone (O3). CO2 is the substrate for photosynthesis and thus can accelerate tree growth, whereas O3 is a highly reactive oxygen species and interferes with basic physiological functions. This review summarizes the impacts of CO2 and O3 on tree chemical composition and highlights the consequences thereof for trophic interactions and ecosystem dynamics. CO2 and O3 influence phytochemical composition by altering substrate availability and biochemical/physiological processes such as photosynthesis and defense signaling pathways. Growth of trees under enriched CO2 generally leads to an increase in the C/N ratio, due to a decline in foliar nitrogen and concomitant increases in carbohydrates and phenolics. Terpenoid levels generally are not affected by atmospheric CO2 concentration. O3 triggers up-regulation of antioxidant defense pathways, leading to the production of simple phenolics and flavonoids (more so in angiosperms than gymnosperms). Tannins levels generally are unaffected, while terpenoids exhibit variable responses. In combination, CO2 and O3 exert both additive and interactive effects on tree chemical composition. CO2-and O3-mediated changes in plant chemistry influence host selection, individual performance (development, growth, reproduction), and population densities of herbivores (primarily phytophagous insects) and soil invertebrates. These changes can effect shifts in the amount and temporal pattern of forest canopy damage and organic substrate deposition. Decomposition rates of leaf litter produced under elevated CO2 and O3 may or may not be altered, and can respond to both the independent and interactive effects of the pollutants. Overall, however, CO2 and O3 effects on decomposition will be influenced more by their impacts on the quantity, rather than quality, of litter produced. A prominent theme to emerge from this and related reviews is that the effects of elevated CO2 and O3 on plant chemistry and ecological interactions are highly context- and species-specific, thus frustrating attempts to identify general, global patterns. Many of the interactions that govern above- and below-ground community and ecosystem processes are chemically mediated, ultimately influencing terrestrial carbon sequestration and feeding back to influence atmospheric composition. Thus, the discipline of chemical ecology is fundamentally important for elucidating the impacts of humans on the health and sustainability of forest ecosystems. Future research should seek to increase the diversity of natural products, species, and biomes studied; incorporate long-term, multi-factor experiments; and employ a comprehensive “genes to ecosystems” perspective that couples genetic/genomic tools with the approaches of evolutionary and ecosystem ecology.

(A.M.T.) Understanding and predicting the impacts of anthropogenically driven climate change on species interactions and ecosystem processes is a critical scientific and societal challenge. Climate change has important ecological consequences for species interactions that occur across multiple trophic levels. In this Update, we broadly examine recent literature focused on disentangling the direct and indirect effects of temperature and water availability on plants, phytophagous insects, and the natural enemies of these insects, with special attention given to forest ecosystems. We highlight the role of temperature in shaping plant and insect metabolism, growth, development, and phenology. Additionally, we address the complexity involved in determining climate-mediated effects on plant-insect and multitrophic level interactions as well as the roles of plant ecophysiological processes in driving both bottom-up and top-down controls. Climate warming may exacerbate plant susceptibility to attack by some insect groups, particularly under reduced water availability. Despite considerable growth in research investigating the effects of climate change on plants and insects, we lack a mechanistic understanding of how temperature and precipitation influence species interactions, particularly with respect to plant defense traits and insect outbreaks. Moreover, a systematic literature review reveals that research efforts to date are highly overrepresented by plant studies and suggests a need for greater attention to plant-insect and multitrophic level interactions. Understanding the role of climatic variability and change on such interactions will provide further insight into links between abiotic and biotic drivers of community-and ecosystem-level processes.Anthropogenic activities have led to rapid and unprecedented increases in atmospheric carbon dioxide (CO 2 ) and other greenhouse gases, which in turn have resulted in numerous observable climatic changes, such as elevated temperature, increased frequency and severity of extreme weather events (e.g. heat waves and droughts), and altered precipitation patterns (e.g. decreased snow cover) (National Research Council, 2010). Species are responding to these climate change factors, as demonstrated by shifts in phenology (the timing of key biological and life history events), biogeographic ranges, and ecological interactions (Bale et al., 2002;Parmesan and Yohe, 2003;Hegland et al., 2009;Robinson et al., 2012). In this Update, we review and discuss the consequences of climate change on plant-insect and multitrophic interactions. Specifically, we address the direct and indirect effects of climate warming and altered precipitation patterns on plants, phytophagous insects, and higher trophic level organisms. We focus on these two components of climate change, firstly, because temperature is the abiotic factor that most directly influences insects (Bale et al., 2002), and secondly, because water availability plays a prominent role in mediating plant-insect interactions (Mattso...

Summary1. The impacts of elevated atmospheric CO 2 and/or O 3 have been examined over 4 years using an open-air exposure system in an aggrading northern temperate forest containing two different functional groups (the indeterminate, pioneer, O 3 -sensitive species Trembling Aspen, Populus tremuloides and Paper Birch, Betula papyrifera , and the determinate, late successional, O 3 -tolerant species Sugar Maple, Acer saccharum ). 2. The responses to these interacting greenhouse gases have been remarkably consistent in pure Aspen stands and in mixed Aspen/Birch and Aspen/Maple stands, from leaf to ecosystem level, for O 3 -tolerant as well as O 3 -sensitive genotypes and across various trophic levels. These two gases act in opposing ways, and even at low concentrations (1·5 × ambient, with ambient averaging 34 -36 nL L − 1 during the summer daylight hours), O 3 offsets or moderates the responses induced by elevated CO 2 . 3. After 3 years of exposure to 560 µ mol mol − 1 CO 2 , the above-ground volume of Aspen stands was 40% above those grown at ambient CO 2 , and there was no indication of a diminishing growth trend. In contrast, O 3 at 1·5 × ambient completely offset the growth enhancement by CO 2 , both for O 3 -sensitive and O 3 -tolerant clones. Implications of this finding for carbon sequestration, plantations to reduce excess CO 2 , and global models of forest productivity and climate change are presented.