systemin and ethylene

Topics: Abscisic Acid; Adaptation, Physiological; Cyclopentanes; Cysteine Endopeptidases; Cytokinins; Ethylenes; Mitogen-Activated Protein Kinases; Multienzyme Complexes; Oxylipins; Peptides; Plant Physiological Phenomena; Proteasome Endopeptidase Complex; Signal Transduction; Ubiquitins

The wound response of tomato plants has been extensively studied, and provides a useful model to understand signal transduction events leading from injury to marker gene expression. The principal markers that have been used in these studies are genes encoding proteinase inhibitor (pin) proteins. Activation of pin genes occurs in the wounded leaf and in distant unwounded leaves of the plant. This paper reviews current understanding of signalling pathways in the wounded leaf, and in the systemically responding unwounded leaves. First, the nature of known elicitors and their potential roles in planta are discussed, in particular, oligogalacturonides, jasmonates and the peptide signal, systemin. Inhibitors of wound-induced proteinase inhibitor (pin) expression are also reviewed, with particular reference to phenolics, sulphydryl reagents and fusicoccin. In each section, results obtained from the bioassay are considered within the wider context of data from mutants and from transgenic plants with altered levels of putative signalling components. Following this introduction, current models for pin gene regulation are described and discussed, together with a summary for the involvement of phosphorylation-dephosphorylation in wound signalling. Finally, a new model for wound-induced pin gene expression is presented, arising from recent data from the author's laboratory.

Topics: Amino Acid Sequence; Cyclopentanes; Ethylenes; Gene Expression Regulation, Plant; Lipid Metabolism; Models, Biological; Molecular Sequence Data; Mutation; Oxylipins; Peptides; Plants, Genetically Modified; Polysaccharides; Protease Inhibitors; Signal Transduction; Solanum lycopersicum

Topics: Acetates; Cyclopentanes; Ethylenes; Immunity, Innate; Oxylipins; Peptides; Plant Growth Regulators; Plant Proteins; Plants; Salicylates; Salicylic Acid; Signal Transduction

Systemins and their hydroxyproline-rich glycopeptide systemin (ppHS) subfamily members are known to mediate antiherbivore defenses in some solanaceous taxa but not others; functions other than in defense remain largely unexplored. Nicotiana attenuata's ppHS is known not to function in herbivore defense. NappHS transcripts are abundant in flowers, particularly in pistils, and when two N. attenuata accessions from Utah and Arizona were transformed to silence NappHS by RNAi (IRsys), seed capsule production and seed number per capsule were reduced in both accessions. These reductions in reproductive performance could not be attributed to impaired pollen or ovule viability; hand-pollination of all IRsys lines of both accessions restored seed production per capsule to levels found in wild-type plants. Rather, changes in flower morphology that decreased the efficiency of self-pollination are likely responsible: IRsys plants of both accessions have flowers with pistils that protrude beyond their anthers. Because these changes in flower morphology are reminiscent of CORONATINE-INSENSITIVE1-silenced N. attenuata plants, we measured jasmonates (JAs) and their biosynthetic transcripts in different floral developmental stages, and found levels of JA-isoleucine (Ile)/leucine and threonine deaminase transcripts, which are abundant in wild-type pistils, to be significantly reduced in IRsys buds and flowers. Threonine deaminase supplies Ile for JA-Ile biosynthesis, and we propose that ppHS mediates JA signaling during flower development and thereby changes flower morphology. These results suggest that the function of ppHS family members in N. attenuata may have diversified to modulate flower morphology and thereby outcrossing rates in response to biotic or abiotic stresses.

Topics: Abscisic Acid; Arizona; Cyclopentanes; Ethylenes; Feeding Behavior; Flowers; Gene Expression Profiling; Gene Expression Regulation, Plant; Gene Silencing; Genes, Plant; Glycopeptides; Hydroxyproline; Indoleacetic Acids; Nicotiana; Oxylipins; Peptides; Pollination; Protein Precursors; RNA, Messenger; Seeds; Utah

To determine whether the Nicotiana tabacum preproHypSys homolog in Nicotiana attenuata (NapreproHypSys) mediates anti-herbivore responses, we silenced (IRsys) and ectopically over-expressed (OVsys) NapreproHypSys in N. attenuata. Neither herbivore simulation nor methyl jasmonate (MeJA) application increased transcripts in wild-type (WT) or transformed lines. Compared to WT plants, OVsys plants had marginally higher constitutive levels but normally induced levels of trypsin proteinase inhibitors (TPIs) and nicotine; IRsys plants did not differ from WT plants. Herbivory-associated signalling [salicylic acid-induced protein kinase (SIPK) activity, jasmonic acid (JA), jasmonic acid-isoleucine/leucine (JA-Ile/Leu) and ethylene production or perception] did not differ strongly among the lines, but JA, JA-Ile/Leu and ethylene were marginally higher in OVsys plants. Manduca sexta larval performance did not differ among the lines, but feeding induced levels of TPI and nicotine in OVsys plants and decreased them in IRsys plants relative to WT. The secondary metabolite profiles of plants transplanted into N. attenuata's native habitat in the Great Basin Desert (UT, USA) mirrored those of glasshouse-grown plants, and compared to WT plants, OVsys plants suffered marginally less damage from grasshoppers, mirids and flea beetles but did not differ in their ability to attract Geocoris predators. We conclude that NapreproHypSys does not play a central role in anti-herbivore defense signalling in this native tobacco.

Topics: Acetates; Animals; Cyclopentanes; Ethylenes; Feeding Behavior; Gene Expression Regulation, Plant; Glycopeptides; Hydroxyproline; Larva; Manduca; Nicotiana; Oxylipins; Peptides; Plants, Genetically Modified; RNA, Messenger; Signal Transduction