Plant interactions with microbes and insects: from molecular mechanisms to ecology

Plants are members of complex communities and interact both with antagonists and beneficial organisms. An important question in plant defense-signaling research is how plants integrate signals induced by pathogens, beneficial microbes and insects into the most appropriate adaptive response. Molecular and genomic tools are now being used to uncover the complexity of the induced defense signaling networks that have evolved during the arms races between plants and their attackers. Molecular biologists and ecologists are joining forces to place molecular mechanisms of plant defense into an ecological perspective. Here, we review our current understanding of the molecular mechanisms of induced plant defense and their potential ecological relevance in nature.     

生物和非生物逆境胁迫下的植物系统信号

复杂多变的自然环境使植物进化出许多适应策略, 其中由局部胁迫引起的系统响应广泛存在, 精细调节植物的生长发育和环境适应能力。植物系统响应的诱导因素首先引起植物从局部到全株范围的信号转导, 这类信号称为系统信号。当受到外界刺激时, 植物首先在受刺激细胞内触发化学信号分子的变化, 如茉莉酸和水杨酸甲酯等在浓度和信号强度方面发生变化; 进而, 伴随着一系列复杂的信号转换, 多种信号组分共同完成系统响应的激活。植物激素、小分子肽和RNA等被认为是缓慢系统信号通路中的关键组分, 而目前也有大量研究阐释了由活性氧、钙信号和电信号相互偶联组成的快速系统信号通路。植物系统信号对其生存和繁衍至关重要, 其精确的转导机制仍值得深入研究。该文综述了植物响应环境的系统信号转导研究进展, 对关键的系统信号组分及其转导机制进行了总结, 同时对植物系统信号传递的研究方向进行了展望。     

Plant immune system: Basal immunity

Plants have an efficient system of innate immunity that is based on the effective detection of potentially harmful microorganisms and rapid induction of defense responses. The first level of plant immunity is basal immunity, which is induced by the conserved molecular structures of microbes, such as bacterial flagellins or fungal chitin, or molecules that result from the interaction of plants with pathogens, for example oligosaccharides and peptides (“danger signals”). Plants recognize these inducers through receptors localized to the plasma membrane, represented mainly by receptor-like protein kinases or receptor-like proteins. Activation of the receptor by a ligand triggers a complex network of signaling events, which eventually cause an array of plant defense responses to prevent further spread of the pathogen.     

Plant volatiles as method of communication

Plants emit volatile compounds that can act as a communication method to insects, neighboring plants and pathogens. Plants respond to leaf and root damage by herbivores and pathogens by emitting these compounds. The volatile compounds can deter the herbivores or pathogens directly or indirectly by attracting their natural enemies to kill them. The simultaneous damage of plants by herbivores and pathogens can influence plant defense. The induced plant volatiles can also make neighboring plants ready for defense or induce defense in parts distant from the damaged area of the same plant. Belowground root herbivory can alter the defense response to aboveground leaf herbivory. In addition, most plants normally emit volatile compounds from their flowers that directly attract foraging mutualistic insects for nectar, which in turn perform the very important function of pollination for subsequent reproduction. The volatile compounds emitted from the floral and vegetative parts of plants belong to three main classes of compounds: terpenoids, phenylpropanoids/benzenoids, and C6-aldehydes (green-leaf volatiles). The volatile phytohormones methyl salicylate and methyl jasmonate serve as important signaling molecules for communication purposes, and interact with each other to optimize the plant defense response. Here we discuss and integrate the current knowledge on all types of communication between plants and insects, neighboring plants and pathogens that are mediated through plant volatiles.     

Receptor-like kinases and receptor-like proteins: keys to pathogen recognition and defense signaling in plant innate immunity

Plants have evolved multiple layers of defense against various pathogens in the environment. Receptor-like kinases/proteins (RLKs/RLPs) are on the front lines of the battle between plants and pathogens since they are present at the plasma membrane and perceive signature molecules from either the invading pathogen or damaged plant tissue. With a few notable exceptions, most RLKs/RLPs are positive regulators of plant innate immunity. In this review, we summarize recently discovered RLKs/RLPs that are involved in plant defense responses against various classes of pathogens.We also describe what is currently known about the mechanisms of RLK-mediated initiation of signaling via protein-protein interactions and phosphorylation.     

The gene encoding glutathione-dependent formaldehyde dehydrogenase/GSNO reductase is responsive to wounding,jasmonic acid and salicylic acid

It has recently been discovered that glutathione-dependent formaldehyde dehydrogenase (FALDH) exhibits a strong S-nitrosoglutathione reductase activity. Plants use NO and S-nitrosothiols as signaling molecules to activate defense mechanisms. Therefore, it is interesting to investigate the regulation of FALDH by mechanical wounding and plant hormones involved in signal transduction. Our results show that the gene encoding FALDH in Arabidopsis (ADH2) is down-regulated by wounding and activated by salicylic acid (SA). In tobacco, FALDH levels and enzymatic activity decreased after jasmonate treatment, and increased in response to SA. This is the first time that regulation of FALDH in response to signals associated with plant defense has been demonstrated.     

Short- and long-distance signaling in plant defense

When encountering microbial pathogens, plant cells can recognize danger signals derived from pathogens, activate plant immune responses and generate cell-autonomous as well as non-cell-autonomous defense signaling molecules, which promotes defense responses at the infection site and in the neighboring cells. Meanwhile, local damages can result in the release of immunogenic signals including damage-associated molecule patterns and phytocytokines, which also serve as danger signals to potentiate immune responses in cells surrounding the infection site. Activation of local defense responses further induces the production of long-distance defense signals, which can move to distal tissue to activate systemic acquired resistance. In this review, we summarize current knowledge on various signaling molecules involved in short- and long-distance defense signaling, and emphasize the roles of regulatory proteins involved in the processes.     

The stomata frontline of plant interaction with the environment-perspectives from hormone regulation

Plants have evolved elaborate mechanisms to perceive and integrate signals from various environmental conditions.On leaf surface,stomata formed by pairs of guard cells mediate gas exchange,water transp...     

Resistance and Susceptibility of Plants to Fungal Pathogens

Plants are under continuous threat of infection by pathogens endowed with diverse strategies to colonize their host. Comprehensive biochemical and genetic approaches are now starting to reveal the complex signaling pathways that mediate plant disease resistance. Initiation of defense signaling often involves specific recognition of invading pathogens by the products of specialized host resistance (R) genes. Potential resistance signaling components have been identified by mutational analyses to be required for specific resistance in the model Arabidopsis and some crop species. Strikingly, many of the components share similarity to that of innate immune systems in animals. Evidence is also accumulating that plant pathogens have a number of ways to evade host defenses during the early stages of infection, similar to animal pathogens. These strategies are becoming much better understood in a number of plant–pathogen interactions. In this review, we focus on the current knowledge of host factors that control plant resistance and susceptibility to fungal pathogens. The knowledge accumulated in these studies will serve a fundamental basis for combating diseases in strategic molecular agriculture.     

Signaling in plant disease resistance and symbiosis

Interactions between plants and microbes result in plant disease and symbiosis. The former causes considerable economic damage in modern agriculture, while the latter has produced great beneficial effects to our agriculture system. Comparison of the two interactions has revealed that a common panel of signaling pathways might participate in the establishment of the equilibrium between plant and microbes or its break-up. Plants appear to detect both pathogenic and symbiotic microbes by a similar set of genes. All symbiotic microbes seem to produce effectors to overcome plant basal defenses and it is speculated that symbiotic effectors have functions similar to pathogenic ones. Signaling molecules, salicylic acid (SA),jasmonic acid (JA) and ethylene (ET), are involved in both plant defense and symbiosis. Switching off signals contributing to deterioration of disease symptom would establish a new equilibrium between plant and pathogenic microbes. This would facilitate the development of strategies for durable disease resistance.     

CML9, a multifunctional Arabidopsis thaliana calmodulin-like protein involved in stress responses and plant growth?

Plants have evolved complex signaling networks to respond to their fluctuating environment and adapt their growth and development. Calcium-dependent signaling pathways play key role in the onset of these adaptive responses. In plant cells, the intracellular calcium transients are triggered by numerous stimuli and it is supposed that the large repertory of calcium sensors present in higher plants could contribute to integrate these signals in physiological responses. Here, we present data on CML9, a calmodulin-like protein that appears to be involved in plant responses to both biotic and abiotic stress. Using a reverse genetic approach based on gain and loss of function mutants, we present here data indicating that this CML might also be involved in root growth control in response to the flagellin, a pathogen-associated molecular pattern (PAMP) also involved in plant immunity.     

Light signaling and UV‐B‐mediated plant growth regulation

Light plays an important role in plants’ growth and development throughout their life cycle. Plants alter their morphological features in response to light cues of varying intensity and quality. Dedicated photoreceptors help plants to perceive light signals of different wavelengths. Activated photoreceptors stimulate the downstream signaling cascades that lead to extensive gene expression changes responsible for physiological and developmental responses. Proteins such as ELONGATED HYPOCOTYL5 (HY5) and CONSTITUTIVELY PHOTOMORPHOGENIC 1 (COP1) act as important factors which modulate light‐regulated gene expression, especially during seedling development. These factors function as central regulatory intermediates not only in red, far‐red, and blue light pathways but also in the UV‐B signaling pathway. UV‐B radiation makes up only a minor fraction of sunlight, yet it imparts many positive and negative effects on plant growth. Studies on UV‐B perception, signaling, and response in plants has considerably surged in recent times. Plants have developed different strategies to use UV‐B as a developmental cue as well as to withstand high doses of UV‐B radiation. Plants’ responses to UV‐B are an integration of its cross‐talks with both environmental factors and phytohormones. This review outlines the current developments in light signaling with a major focus on UV‐B‐mediated plant growth regulation.     

Signal Molecules for Plant Defense Responses to Biotic Stress

Plants are capable of recognizing the penetrating pathogens and of responding to their attack by the activation of the defense systems. Signal transduction from the receptor to the cell genome is required for this activation. Recently, signal molecules have been found, which are involved in the signal transduction triggered in response to biotic stress. The data accumulated imply the presence of a complex and well-coordinated signal network in plant cells. This net controls plant defense responses to pathogen attacks.     

Cross talk between signaling pathways in pathogen defense

Plant defense in response to microbial attack is regulated through a complex network of signaling pathways that involve three signaling molecules: salicylic acid (SA), jasmonic acid (JA) and ethylene. The SA and JA signaling pathways are mutually antagonistic. This regulatory cross talk may have evolved to allow plants to fine-tune the induction of their defenses in response to different plant pathogens.     

Enzyme Action in the Regulation of Plant Hormone Responses

Plants synthesize a chemically diverse range of hormones that regulate growth, development, and responses to environmental stresses. The major classes of plant hormones are specialized metabolites with exquisitely tailored perception and signaling systems, but equally important are the enzymes that control the dose and exposure to the bioactive forms of these molecules. Here, we review new insights into the role of enzyme families, including the SABATH methyltransferases, the methylesterases, the GH3 acyl acid-amido synthetases, and the hormone peptidyl hydrolases, in controlling the biosynthesis and modifications of plant hormones and how these enzymes contribute to the network of chemical signals responsible for plant growth, development, and environmental adaptation.     

Jasmonate-induced defenses: a tale of intelligence, collaborators and rascals

Plants have sophisticated defense systems to protect their tissues against the attack of herbivorous organisms. Many of these defenses are orchestrated by the oxylipin jasmonate. A growing body of evidence indicates that the expression of jasmonate-induced responses is tightly regulated by the ecological context of the plant. Ecological information is provided by molecular signals that indicate the nature of the attacker, the value of the attacked organs, phytochrome status and thereby proximity of competing plants, association with beneficial organisms and history of plant interactions with pathogens and herbivores. This review discusses recent advances in this field and highlights the need to map the activities of informational modulators to specific control points within our emerging model of jasmonate signaling.     

Shedding light on large-scale chromatin reorganization in Arabidopsis thaliana

Plants need to respond quickly and appropriately to various types of light signals from the environment to optimize growth and development. The immediate response to shading, reduced photon flux (low light), and changes in spectral quality involves changes in gene regulation. In the case of more persistent shade, the plant shows a dramatic change in the organization of chromatin. Both plant responses are controlled via photoreceptor signaling proteins. Recently, several studies have revealed similar features of chromatin reorganization in response to various abiotic and biotic signals, while others have unveiled intricate molecular networks of light signaling towards gene regulation. This opinion paper briefly describes the chromatin (de)compaction response from a light-signaling perspective to provide a link between chromatin and the molecular network of photoreceptors and E3 ubiquitin ligase complexes.