Salicylic acid beyond defence: its role in plant growth and development

In recent years salicylic acid (SA) has been the focus of intensive research due to its function as an endogenous signal mediating local and systemic plant defence responses against pathogens. It has also been found that SA plays a role during the plant response to abiotic stresses such as drought, chilling, heavy metal toxicity, heat, and osmotic stress. In this sense, SA appears to be, just like in mammals, an 'effective therapeutic agent' for plants. Besides this function during biotic and abiotic stress, SA plays a crucial role in the regulation of physiological and biochemical processes during the entire lifespan of the plant. The discovery of its targets and the understanding of its molecular modes of action in physiological processes could help in the dissection of the complex SA signalling network, confirming its important role in both plant health and disease. Here, the evidence that supports the role of SA during plant growth and development is reviewed by comparing experiments performed by exogenous application of SA with analysis of genotypes affected by SA levels and/or perception.     

Salicylic acid and the plant pathogen Erwinia carotovora induce defense genes via antagonistic pathways

Infection of tobacco plants with the plant pathogenic bacterium Erwinia carotovora subsp. carotovora or treatment of plants with Erwinia -derived elicitor preparations leads to the induction of a number of genes thought to play a role in plant defense response to pathogens. In order to determine the role of salicylic acid (SA) in the induction of the Erwinia responsive genes, the accumulation of mRNAs for these and other genes encoding pathogenesis-related proteins (PR genes) in response to both Erwinia elicitors and SA was determined. PR genes were identified which were preferentially induced by Erwinia elicitor preparations, one gene was induced by SA but not by Erwinia , and another gene was induced by both type of treatments. The differential expression of these genes and the timing of induction suggest that SA is not the signal molecule leading to the early response of plants to Erwinia . This was demonstrated by experiments using transgenic NahG plants that overproduce a salicylate hydroxylase inactivating SA. The elicitation of PR genes by Erwinia was similar in NahG and wild-type plants. Therefore, induction of plant defense genes by Erwinia and SA seems to be by two distinct pathways leading to expression of separate sets of genes. Furthermore, we could demonstrate that Erwinia elicitors antagonize the SA-mediated induction of PR genes. Similarly, SA appeared to inhibit the induction of PR genes elicited by Erwinia . The observed antagonism between the two signal transduction pathways indicates the presence of a common regulatory element in both pathways that acts downstream of SA in the SA-mediated response.     

Genome-wide analysis of phenylpropanoid defence pathways

Phenylpropanoids can function as preformed and inducible antimicrobial compounds, as well as signal molecules, in plant-microbe interactions. Since we last reviewed the field 8 years ago, there has been a huge increase in our understanding of the genes of phenylpropanoid biosynthesis and their regulation, brought about largely by advances in genome technology, from whole-genome sequencing to massively parallel gene expression profiling. Here, we present an overview of the biosynthesis and roles of phenylpropanoids in plant defence, together with an analysis of confirmed and predicted phenylpropanoid pathway genes in the sequenced genomes of 11 plant species. Examples are provided of phylogenetic and expression clustering analyses, and the large body of underlying genomic data is provided through a website accessible from the article.     

Importance of plant‐ and microbe‐driven metabolic pathways for plant defence

Expression of plant phenotypes can depend on both plant genomes and interactions between plants and the microbes living in, on and near their roots. We understand a growing number of the mechanistic links between plant genotypes and phenotypes, such as defence against herbivory (see brief review in Hubbard et al., 2019), yet the links between root microbiomes and the comprehensive swathe of plant phenotypes they affect (Friesen et al., 2011) remain less clear. In this issue of Molecular Ecology, Hubbard et al. (2019) follow microbe‐ and plant‐driven changes in plant defence against hervibory from molecular underpinnings to ecological consequences, contrasting both the metabolites affected and the magnitude of defensive impact. Naively, we might expect plant genomes to drive more variation in phenotype than the root microbiome, but Hubbard et al. (2019) find the opposite, implying profound consequences for plant trait evolution and ecological interactions.     

Phospholipid signalling in plant defence

Phospholipid-derived molecules are emerging as novel second messengers in plant defence signalling. Recent research has begun to reveal the signals produced by the enzymes phospholipase C, phospholipase D and phospholipase A2 and their putative downstream targets. These include the activation of a MAP kinase cascade and triggering of an oxidative burst by phosphatidic acid; the regulation of ion channels and proton pumps by lysophospholipids and free fatty acids; and the conversion of free fatty acids into bioactive octadecanoids such as jasmonic acid.     

Long-distance signalling in plant defence

Plants use inducible defence mechanisms to fend off harmful organisms. Resistance that is induced in response to local attack is often expressed systemically, that is, in organs that are not yet damaged. In the search for translocated defence signals, biochemical studies follow the physical movement of putative signals, and grafting experiments use mutants that are impaired in the production or perception of these signals. Long-distance signals can directly activate defence or can prime for the stronger and faster induction of defence. Historically, research has focused on the vascular transport of signalling metabolites, but volatiles can play a crucial role as well. We compare the advantages and constraints of vascular and airborne signals for the plant, and discuss how they can act in synergy to achieve optimised resistance in distal plant parts.     

Role of plant hormones in plant defence responses

Plant hormones play important roles in regulating developmental processes and signaling networks involved in plant responses to a wide range of biotic and abiotic stresses. Significant progress has been made in identifying the key components and understanding the role of salicylic acid (SA), jasmonates (JA) and ethylene (ET) in plant responses to biotic stresses. Recent studies indicate that other hormones such as abscisic acid (ABA), auxin, gibberellic acid (GA), cytokinin (CK), brassinosteroids (BR) and peptide hormones are also implicated in plant defence signaling pathways but their role in plant defence is less well studied. Here, we review recent advances made in understanding the role of these hormones in modulating plant defence responses against various diseases and pests.     

Proteases in pathogenesis and plant defence

Plant pathogens deliver a variety of virulence factors to host cells to suppress basal defence responses and create suitable environments for their propagation. Plants have in turn evolved disease resistance genes whose products detect the virulence factors as a signal of invasion and activate effective defence responses. Understanding how a virulence effector contributes to virulence on susceptible hosts but becomes an avirulence factor that triggers defence responses on resistance hosts has been a major focus in plant research. Recent studies have shown that a growing list of pathogen-encoded effectors functions as proteases that are secreted into plant cells to modify host proteins. In addition, several plant proteases have been found to function in activation of the defence mechanism. These findings reveal that post-translational modification of host proteins through proteolytic processing is a widely used mechanism in regulating the plant defence response.     

A herbivore that manipulates plant defence

Phytopathogens and herbivores induce plant defences. Whereas there is evidence that some pathogens suppress these defences by interfering with signalling pathways involved in the defence, such evidence is scarce for herbivores. We found that the invasive spider mite Tetranychus evansi suppresses the induction of the salicylic acid and jasmonic acid signalling routes involved in induced plant defences in tomato. This was reflected in the levels of inducible defence compounds, such as proteinase inhibitors, which in mite-infested plants were reduced to even lower levels than the constitutive levels in herbivore-free plants. Additionally, the spider mite suppressed the release of inducible volatiles, which are implicated in plant defence. Consequently, the mites performed much better on previously attacked plants than on non-attacked plants. These findings provide a new perspective on plant-herbivore interactions, plant protection and plant resistance to invasive species.     

Salicylic acid regulates flowering time and links defence responses and reproductive development

Flowering relies on signaling networks that integrate endogenous and external cues. Normally, plants flower at a particular season, reflecting day length and/or temperature cues. However, plants can surpass this seasonal regulation and show precocious flowering under stress environmental conditions. Here, we show that UV-C light stress activates the transition to flowering in Arabidopsis thaliana through salicylic acid (SA). Moreover, SA also regulates flowering time in non-stressed plants, as SA-deficient plants are late flowering. The regulation of flowering time by SA seems to involve the photoperiod and autonomous pathways, but it does not require the function of the flowering time genes CONSTANS (CO), FCA, or FLOWERING LOCUS C (FLC).     

Light perception in plant disease defence signalling

Light is a predominant factor in the control of plant growth, development and stress responses. Many biotic stress responses in plants are therefore specifically adjusted by the prevailing light conditions. The plant cell is equipped with sophisticated light-sensing mechanisms that are localised inside and outside of the chloroplast and the nucleus. Recent progress has provided models of how the signalling pathways that are involved in light perception and in defence could operate and interact to form a plant defence network. Such a signalling network includes systems to sense light and regulate gene expression. Photo-produced H(2)O(2) and other reactive oxygen species in the cell also play an essential role in this regulatory network, controlling biotic and abiotic stress responses.     

The role of lectins in plant defence

Summary Recent progress in the search for the physiological role of plant lectins supports the idea that some of these proteins are involved in the defence mechanisms of the plant. To place the evidence in favour of such a defensive role in a broad perspective, a short overview is given of the most important plant pathogens and predators. In addition, the solutions that plants have developed to resist the continuous threat of a hostile environment are briefly discussed in relation to the protective role of proteins in general. The presumed involvement of plant lectins in defence mechanisms is first inferred from an analysis of the biochemical, physiological, cellular biological and molecular biological properties of plant lectins. Subsequently, the available experimental evidence for the involvement of lectins in the plant's defence against viruses, bacteria, fungi and herbivorous invertebrates and vertebrates is discussed in some detail. Since the defensive role of plant lectins is determined largely by their ability to recognize and bind foreign glycans, a brief discussion is given of how the basically protective properties of these proteins can be exploited for histochemical applications in biological and biomedical research.