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Manipulations that alter microbial communities of colonized plants can be used to detect changes in plant immune systems, including the activation of PTI and ETI. Here we assess the functional redundancies of immune-stimulating bacterial complexes (ISBCs) that consist of flg22, elf18, PGN, and cef18, a variant of EF-Tu, in the activation of immune response in plants. We find that immune responses triggered by ISBCs have distinct patterns of functional redundancy. The observed variation in SGI in response to ISBCs is explained by plant variation in PTI, genetic variation in ETI, and a possible MAMP-specific fine-tuning of the immune response. Collectively, the data suggest that the plant immune system has evolved to detect signals that are relevant to the microbe, including MAMPs. By functionally uniting previous MAMP findings with each other and to the host microbial community, this work provides a framework for interpretation of diverse MAMP signals.

Recent studies indicate that disease and colonization by a single beneficial microbe can have different effects on plant immunity [ 10 ]. However, whether these responses are common throughout the plant kingdom remains unknown. In this study, we report that the perception of two MAMPs and one bacterial factor can differentially affect SGI. We also describe the existence of novel components that modulate MAMP perception. This detailed quantitative analysis of MAMP perception in multiple plant species will be valuable for comparative studies of immunity to infection and can help inform the design of biotechnologies that maximize the value of beneficial microbes.

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MAMP elicited MTI (elicitor-triggered immunity, also called PAMP triggered immunity or PTI), is the most commonly studied branch of plant immunity. It is important in the defense against pathogens and promotes basal levels of immunity against non-pathogenic microorganisms [ 17 ]. This defense system is readily perceivable by loss-of-function mutants of regulatory genes in plants, which are compromised in their capacity to perceive and respond to microbe-associated elicitors. Recent investigations of the molecular network and its crosstalk with abiotic stress responses have established that it is a complex and tightly integrated network that operates upstream of abiotic stress signaling to activate general and basal immunity to pathogens and abiotic stress-inducing agents [ 22 ]. Plants have evolved sensing mechanisms that recognize specific microbe-associated elicitors by specialized molecular sensors (also called pattern recognition receptors, or PRRs) localized in the plasma membrane or internal membranes [ 3, 29, 30 ]. They include 13 receptors to EF-Tu [ 26, 32 ], 24 receptors to Flg [ 27, 33 ], 18 receptors to PGN [ 20, 34 ], 14 receptors to chitin [ 28 ], and nine receptors to -glucans [ 18, 31 ]. Receptor activation leads to a signal transduction cascade that is ultimately monitored by many negative regulators, which fine-tune immune responses. These signals include calcium waves, reactive oxygen species (ROS), MAP kinases and other regulators [ 27 ]. Finally, ROS and MAP kinase cascades induce a downstream transcriptional program, which includes the expression of defence-related genes and the production of antimicrobial compounds [ 17, 35 ]. Thus, a successful plant pathogen must overcome these layers of protection before reaching its host [ 35 ]. One strategy for pathogen attack is to suppress PTI by employing MAMPs [ 3, 24 ]. An example is the effect of the MAMP flagellin in fungal biotrophy [ 36 ]. ABA and calcium are known to play an important role in triggering MTI by interacting with other signalling components, among which are calcium dependent protein kinases (CDPKs) and MAP kinases [ 36 ]. Thus, it appears that different MAMPs trigger MTI by distinct mechanisms (e.g. flagellin’s effect on ROS production and calcium waves in Arabidopsis) [ 36 ]. On the other hand, flg22 is known to have overlapping but not identical downstream signalling effects with flg22 from Xanthomonas [ 37 ]. MAMP-elicited MTI provides the plant with the ability to distinguish between potential threats and thus renders plants more resistant to pathogens that attack by unknown mechanisms. In contrast to biotrophs, hemibiotrophs are unable to secrete proteins that suppress PTI. Thus, hemibiotrophs are able to colonize the plant for a prolonged period of time. There is evidence that hemibiotrophs have evolved to evade PTI [ 35 ] and express analogues of effectors that block PRRs [ 38 ]. Another difference between Xanthomonas and Arabidopsis is that the latter has additional layers of immunity, i.e. induced systemic resistance (ISR). In Arabidopsis, MAMP perception, activation of MAPKs and the cross-talk with defence hormones, such as jasmonic acid (JA), ethylene (ET), and salicylic acid (SA), are key steps in the induction of ISR [ 39 ]. Therefore, contrary to the resistance mechanism of PTI, ISR requires long-lasting effects. Eventually, the balance between resistance mechanisms is of critical importance to avoid plant attacks by microbes.

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Flg22 exhibits selective pressure on evolution of its receptor, which implies that changes in the receptor might predispose a plant to receive and respond to certain MAMPs [ 4 ]. Therefore, we compared sequence variation of the toll-like receptor FLS2 in 27 previously analyzed strains of A. thaliana [ 36 ] to sequence variation in three MAMP-insensitive genotypes ( Fig 3a ). In these genotypes, FLS2 sequences showed a total of six amino acid differences between the membrane protein domain of receptor (10 of the 1583 amino acids). The FLS2 sequences of the three genotypes are highly similar ( Fig 3b ). Moreover, 26 of the 27 A. thaliana strains have approximately two or three sequence differences with each other. We tested if this sequence variation in A. thaliana corresponded to sequence variation in the FLS2 receptor of MAMP-insensitive genotypes. We selected eight MAMP-insensitive genotypes, each of which is a genotype of the A. thaliana panel in Atwell et al. [ 21 ]. The A. thaliana panel in Atwell et al. is a collection of 186 genotypes that were previously characterized for elf18-induced SGI in response to P. syringae strain B728a [ 3 ]. We sequenced FLS2 in 16 of the eight genotypes. Three of the 16 genotypes carried a single allele that differed from the reference sequence by two amino acids. Three genotypes carried two alleles differing by two amino acids and two more genotypes carried more than two alleles differing by two amino acids. No genotype carried all three alleles differing by two amino acids. If these polymorphisms were functionally relevant, it would be unlikely to find an MAMP-insensitive genotype that carries all three alleles. Moreover, two of the three genotypes that carry all three alleles have a flg22-induced SGI response that is as strong as the SGI response observed in the 186 genotypes of the Atwell et al. panel.

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What’s new in MAMP PRO 5.0.5.3998

Added functionality for exporting data and image for faster analysis.
Added two new views for faster display
Added views for common MAMP databases
Includes new summarization views for displaying popular MAMP entries based on the total amount of searches during the last three months
Format corrections
Many bug fixes