Genome-wide functional analyses of plant coiled-coil NLR-type pathogen receptors reveal essential roles of their N-terminal domain in oligomerization, networking, and immunity.

The ability to induce a defense response after pathogen attack is a critical feature of the immune system of any organism. Nucleotide-binding leucine-rich repeat receptors (NLRs) are key players in this process and perceive the occurrence of nonself-activities or foreign molecules. In plants, coevolution with a variety of pests and pathogens has resulted in repertoires of several hundred diverse NLRs in single individuals and many more in populations as a whole. However, the mechanism by which defense signaling is triggered by these NLRs in plants is poorly understood. Here, we show that upon pathogen perception, NLRs use their N-terminal domains to transactivate other receptors. Their N-terminal domains homo- and heterodimerize, suggesting that plant NLRs oligomerize upon activation, similar to the vertebrate NLRs; however, consistent with their large number in plants, the complexes are highly heterometric. Also, in contrast to metazoan NLRs, the N-terminus, rather than their centrally located nucleotide-binding (NB) domain, can mediate initial partner selection. The highly redundant network of NLR interactions in plants is proposed to provide resilience to perturbation by pathogens.

Innate immunity: has poplar made its BED?

The perennial plant model species Populus trichocarpa has received considerable attention in the last 5 yr because of its potential use as a bioenergy crop. The completion of its genome sequence revealed extensive homologies with the herbaceous annual species Arabidopsis thaliana. This review highlights the similarities and differences at the qualitative defence response components level, notably in putative NBS-LRR protein content and downstream defence regulators. With almost a twofold NBS-LRR gene complement compared with A. thaliana, P. trichocarpa also encodes some putative R-proteins with unusual architectures and possible DNA-binding capacity. P. trichocarpa also possesses all the known main components characteristic of TIR-NB-LRR and CC-NB-LRR signalling. However, very little has been done with regard to the components involved in the poplar qualitative response to pathogens. In addition, the relationship between plant-biotroph perception/signalling and the role of salicylic acid, an important defence compound, remains uncertain. This review aims to identify the genomic components present in poplar that could potentially participate in the qualitative response and highlights where efforts should be devoted to obtain a better understanding of the poplar qualitative defence response.

[System of innate immunity in plants].

The review deals with the mechanisms of innate immunity in plants with a focus on families of pattern-recognition receptors and regard for recent data on complete sequencing of the genomes of several plant species. Plants utilize several families of such receptors, both membrane-bound and cytoplasmic ones, which contain conservative leucine-rich repeats. The lack of adaptive immunity and there to related rearrangements in genes encoding immune receptors in plants are partly compensated by mechanisms of "specific" immunity to counter particular pathogens; such mechanisms being fully encoded within a plant genome. At the level of intracellular signal transduction, as well as in respect of effector mechanisms, similarities between plant and animal innate immune systems can be found, although the latter has many additional aspects.

Genetic insights into underground responses to Fusarium graminearum infection in wheat.

The ongoing global intensification of wheat production will likely be accompanied by a rising pressure of Fusarium diseases. While utmost attention was given to Fusarium head blight (FHB) belowground plant infections of the pathogen have largely been ignored. The current knowledge about the impact of soil borne Fusarium infection on plant performance and the underlying genetic mechanisms for resistance remain very limited. Here, we present the first large-scale investigation of Fusarium root rot (FRR) resistance using a diverse panel of 215 international wheat lines. We obtained data for a total of 21 resistance-related traits, including large-scale Real-time PCR experiments to quantify fungal spread. Association mapping and subsequent haplotype analyses discovered a number of highly conserved genomic regions associated with resistance, and revealed a significant effect of allele stacking on the stembase discoloration. Resistance alleles were accumulated in European winter wheat germplasm, implying indirect prior selection for improved FRR resistance in elite breeding programs. Our results give first insights into the genetic basis of FRR resistance in wheat and demonstrate how molecular parameters can successfully be explored in genomic prediction. Ongoing work will help to further improve our understanding of the complex interactions of genetic factors influencing FRR resistance.

Genome-wide analysis of the soybean CRK-family and transcriptional regulation by biotic stress signals triggering plant immunity.

Cysteine-rich receptor-like kinases (CRKs) are transmembrane proteins that exhibit ectodomains containing the domain of unknown function 26 (DUF26). The CRKs form a large subfamily of receptor-like kinases in plants, and their possible functions remain to be elucidated. Several lines of evidence suggest that CRKs play important roles in plant defense responses to environmental stress, including plant immunity. We performed a genome-wide analysis of CRK encoding genes in soybean (Glycine max). We found 91 GmCRKs distributed in 16 chromosomes, and identified several tandem and segmental duplications, which influenced the expansion of this gene family. According to our phylogenetic analysis, GmCRKs are grouped in four clades. Furthermore, 12% of the members exhibited GmCRKs with a duplicated bi-modular organization of the ectodomains, containing four DUF26 domains. Expression analysis of GmCRKs was performed by exploring publicly available databases, and by RT-qPCR analysis of selected genes in soybean leaves responding to biotic stress signals. GmCRKs exhibited diverse expression patterns in leaves, stems, roots, and other tissues. Some of them were highly expressed in only one type of tissue, suggesting predominant roles in specific tissues. Furthermore, several GmCRKs were induced with PAMPs, DAMPs and the pathogens Phakopsora pachyrhizi and Phytophthora sojae. Expression profiles of several GmCRKs encoding highly similar proteins exhibited antagonist modes of regulation. The results suggest a fine-tuning control of GmCRKs transcriptional regulation in response to external stimuli, including PAMPs and DAMPs. This study offers a comprehensive view of the GmCRKs family in soybean, and provides a foundation for evolutionary and functional analysis of this family of plant proteins involved in the perception of pathogens and activation of plant immunity.

Arabidopsis LBP/BPI related-1 and -2 bind to LPS directly and regulate PR1 expression.

Lipopolysaccharide (LPS) is a major constituent of the outer membrane of Gram-negative bacteria and acts as a pathogen-associated molecular pattern that triggers immune responses in both plants and animals. LPS-binding protein (LBP) and bactericidal/permeability-increasing protein (BPI), which bind to LPS and play important roles in immunity of mammals, have been well studied. However, the molecule contributing to LPS binding in plants is mostly unknown. The Arabidopsis genome carries two genes encoding LBP/BPI-related proteins which we designated as AtLBP/BPI related-1 (AtLBR-1) and AtLBP/BPI related-2 (AtLBR-2). We found that their N-terminal domains were co-purified with cell wall-derived LPS when expressed in E. coli. Since this finding implied the direct binding of AtLBRs to LPS, we also confirmed binding by using LPS-free AtLBRs and purified LPS. AtLBRs directly bind to both rough and smooth types of LPS. We also demonstrated that LPS-treated atlbr mutant Arabidopsis exhibit a significant delay of induction of defence-related gene pathogenesis-related 1 (PR1) but no other PR genes. Furthermore, LPS-treated atlbr mutants showed defects in reactive oxygen species (ROS) generation. These results demonstrate that, as well as LBP and BPI of mammals, AtLBRs also play an important role in the LPS-induced immune response of plants.