Herbivorous insects use diverse feeding strategies to obtain nutrients from their host plants. Rather than acting as passive victims in these interactions, plants respond to herbivory with the production of toxins and defensive proteins that target physiological processes in the insect. Herbivore-challenged plants also emit volatiles that attract insect predators and bolster resistance to future threats. This highly dynamic form of immunity is initiated by the recognition of insect oral secretions and signals from injured plant cells. These initial cues are transmitted within the plant by signal transduction pathways that include calcium ion fluxes, phosphorylation cascades, and, in particular, the jasmonate pathway, which plays a central and conserved role in promoting resistance to a broad spectrum of insects. A detailed understanding of plant immunity to arthropod herbivores will provide new insights into basic mechanisms of chemical communication and plant-animal coevolution and may also facilitate new approaches to crop protection and improvement.

Plant Immunity to Insect Herbivores

In his introduction to this detailed survey of knowledge of insect societies, the author points out that research on insect sociology has proceeded in three phases, the natural history phase, the physiological phase and the population-biology phase. Advances in the first two phases have permitted embarkation in the third phase on a more rigorous theory of social evolution based on population genetics and writing this book, the author wished to relate the three phases of research on insects and to express insect sociology as population biology. A glossary of terms, a considerable bibliography and a general index are included. Other CABI sites 

The Insect Societies

Plant responses to the projected future levels of CO2 were first characterized in short-term experiments lasting days to weeks. However, longer term acclimation responses to elevated CO2 were subsequently discovered to be very important in determining plant and ecosystem function. Free-Air CO2 Enrichment (FACE) experiments are the culmination of efforts to assess the impact of elevated CO2 on plants over multiple seasons and, in the case of crops, over their entire lifetime. FACE has been used to expose vegetation to elevated concentrations of atmospheric CO2 under completely open-air conditions for nearly two decades. This review describes some of the lessons learned from the long-term investment in these experiments. First, elevated CO2 stimulates photosynthetic carbon gain and net primary production over the long term despite down-regulation of Rubisco activity. Second, elevated CO2 improves nitrogen use efficiency and, third, decreases water use at both the leaf and canopy scale. Fourth, elevated CO2 stimulates dark respiration via a transcriptional reprogramming of metabolism. Fifth, elevated CO2 does not directly stimulate C4 photosynthesis, but can indirectly stimulate carbon gain in times and places of drought. Finally, the stimulation of yield by elevated CO2 in crop species is much smaller than expected. While many of these lessons have been most clearly demonstrated in crop systems, all of the lessons have important implications for natural systems.

Elevated CO2 effects on plant carbon, nitrogen, and water relations: six important lessons from FACE

Plants respond to herbivory through various morphological, biochemicals, and molecular mechanisms to counter/offset the effects of herbivore attack. The biochemical mechanisms of defense against the herbivores are wide-ranging, highly dynamic, and are mediated both by direct and indirect defenses. The defensive compounds are either produced constitutively or in response to plant damage, and affect feeding, growth, and survival of herbivores. In addition, plants also release volatile organic compounds that attract the natural enemies of the herbivores. These strategies either act independently or in conjunction with each other. However, our understanding of these defensive mechanisms is still limited. Induced resistance could be exploited as an important tool for the pest management to minimize the amounts of insecticides used for pest control. Host plant resistance to insects, particularly, induced resistance, can also be manipulated with the use of chemical elicitors of secondary metabolites, which confer resistance to insects. By understanding the mechanisms of induced resistance, we can predict the herbivores that are likely to be affected by induced responses. The elicitors of induced responses can be sprayed on crop plants to build up the natural defense system against damage caused by herbivores. The induced responses can also be engineered genetically, so that the defensive compounds are constitutively produced in plants against are challenged by the herbivory. Induced resistance can be exploited for developing crop cultivars, which readily produce the inducible response upon mild infestation, and can act as one of components of integrated pest management for sustainable crop production.

https://www.tandfonline.com/doi/pdf/10.4161/psb.21663

Mechanisms of plant defense against insect herbivores

Volatile compounds act as a language that plants use for their communication and interaction with the surrounding environment. To date, a total of 1700 volatile compounds have been isolated from more than 90 plant families. These volatiles, released from leaves, flowers, and fruits into the atmosphere and from roots into the soil, defend plants against herbivores and pathogens or provide a reproductive advantage by attracting pollinators and seed dispersers. Plant volatiles constitute about 1% of plant secondary metabolites and are mainly represented by terpenoids, phenylpropanoids/benzenoids, fatty acid derivatives, and amino acid derivatives. In this review we focus on the functions of plant volatiles, their biosynthesis and regulation, and the metabolic engineering of the volatile spectrum, which results in plant defense improvement and changes of scent and aroma properties of flowers and fruits.

Plant Volatiles: Recent Advances and Future Perspectives

When Manduca sexta attacks Nicotiana attenuata, fatty acid-amino acid conjugates (FACs) in the larvae's oral secretions (OS) are introduced into feeding wounds. These FACs trigger a transcriptional response that is similar to the response induced by insect damage. Using two-dimensional gel electrophoresis, matrix-assisted laser desorption ionization-time of flight, and liquid chromatography-tandem mass spectrometry, we characterized the proteins in phenolic extracts and in a nuclear fraction of leaves elicited by larval attack, and/or in leaves wounded and treated with OS, FAC-free OS, and synthetic FACs. Phenolic extracts yielded approximately 600 protein spots, many of which were altered by elicitation, whereas nuclear protein fractions yielded approximately 100 spots, most of which were unchanged by elicitation. Reproducible elicitor-induced changes in 90 spots were characterized. In general, proteins that increased were involved in primary metabolism, defense, and transcriptional and translational regulation; those that decreased were involved in photosynthesis. Like the transcriptional defense responses, proteomic changes were strongly elicited by the FACs in OS. A semiquantitative reverse transcription-PCR approach based on peptide sequences was used to compare transcript and protein accumulation patterns for 17 candidate proteins. In six cases the patterns of elicited transcript accumulation were consistent with those of elicited protein accumulation. Functional analysis of one of the identified proteins involved in photosynthesis, RuBPCase activase, was accomplished by virus-induced gene silencing. Plants with decreased levels of RuBPCase activase protein had reduced photosynthetic rates and RuBPCase activity, and less biomass, responses consistent with those of herbivore-attacked plants. We conclude that the response of the plant's proteome to herbivore elicitation is complex, and integrated transcriptome-proteome-metabolome analysis is required to fully understand this ubiquitous ecological interaction.

http://www.plantphysiol.org/content/plantphysiol/142/4/1621.full.pdf

Molecular Interactions between the Specialist Herbivore Manduca sexta (Lepidoptera, Sphingidae) and Its Natural Host Nicotiana attenuata. VII. Changes in the Plant's Proteome

To determine if damage to foliage by biotic agents, including arthropods, fungi, bacteria and viral pathogens, universally downregulates the expression of genes involved in photosynthesis, we compared transcriptome data from microarray experiments after twenty two different forms of biotic damage on eight different plant species. Transcript levels of photosynthesis light reaction, carbon reduction cycle and pigment synthesis genes decreased regardless of the type of biotic attack. The corresponding upregulation of genes coding for the synthesis of jasmonic acid and those involved in the responses to salicylic acid and ethylene suggest that the downregulation of photosynthesis-related genes was part of a defence response. Analysis of the sub-cellular targeting of co-expressed gene clusters revealed that the transcript levels of 84% of the genes that carry a chloroplast targeting peptide sequence decreased. The majority of these downregulated genes shared common regulatory elements, such as G-box (CACGTG), T-box (ACTTTG) and SORLIP (GCCAC) motifs. Strong convergence in the response of transcription suggests that the universal downregulation of photosynthesis-related gene expression is an adaptive response to biotic attack. We hypothesize that slow turnover of many photosynthetic proteins allows plants to invest resources in immediate defence needs without debilitating near term losses in photosynthetic capacity.

https://www.onlinelibrary.wiley.com/doi/pdf/10.1111/j.1365-3040.2010.02167.x

Biotic stress globally downregulates photosynthesis genes.

Elevated CO2 and temperature are altering the interactions between plants and insects with important implications for food security and natural ecosystems. Ecologically, the acceleration of plant phenology by warming is generating mismatches between plants and insect pollinators. Similarly, shifting

/pdf/climate-change-resetting-plant-insect-interactions-r0xbf7ednw.pdf

Climate change: Resetting plant-insect interactions

Evidence for the in planta defensive function of trypsin protease inhibitors (TPIs) comes from observations of enhanced herbivore resistance after heterologous TPI expression or the manipulation of signal cascades that activate numerous defense responses, including TPI production; no studies have altered the expression of an endogenous pi gene to examine defensive function. We isolated two genes with seven- and six-repeat TPI domains from Nicotiana attenuata from the potato (Solanum tuberosum) PI-II family. To determine whether endogenous TPIs in N. attenuata function defensively against the native herbivores, hornworm (Manduca sexta) and mirids (Tupiocoris notatus), we expressed 175 bp of the seven-domain pi from N. attenuata in an antisense orientation in a TPI-producing genotype to reduce TPI expression and expressed the full-length seven-domain pi in a sense orientation under control of a constitutive promoter to restore TPI activity in a natural genotype from Arizona unable to produce TPIs. Constitutive and inducible TPI production in two antisense lines were diminished by 80% to 90% and 33% to 52%, respectively, and sense expression restored 67% of the activity found in the TPI-producing genotype after caterpillar attack in the TPI-deficient A genotype. Hornworm larvae fed on genotypes with low or no TPI activity grew faster, had higher survivorship, and produced heavier pupae than those that fed on genotypes with high TPI activity. T. notatus showed higher preference for genotypes with low or no TPI activity than for genotypes with high TPI levels. We conclude that endogenous TPIs are an effective defense against these native herbivores.

http://www.plantphysiol.org/content/plantphysiol/134/3/1181.full.pdf

Manipulation of endogenous trypsin proteinase inhibitor production in Nicotiana attenuata demonstrates their function as antiherbivore defenses

https://www.life.illinois.edu/delucia/PUBLICATIONS/Nabity%202008.pdf

Jorge A. Zavala

Papers

Molecular Interactions between the Specialist Herbivore Manduca sexta (Lepidoptera, Sphingidae) and Its Natural Host Nicotiana attenuata. VII. Changes in the Plant's Proteome

Biotic stress globally downregulates photosynthesis genes.

Climate change: Resetting plant-insect interactions

Manipulation of endogenous trypsin proteinase inhibitor production in Nicotiana attenuata demonstrates their function as antiherbivore defenses

Indirect suppression of photosynthesis on individual leaves by arthropod herbivory.