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It was recently shown that numerous micro RNAs target nucleotide sequences encoding conserved motifs of NLRs (e.g., P-loop) in many flowering plants (21). The distribution of the individual domains constitutive of NLRs (NB-ARC, NACHT, and LRR) and the different groups of NLRs are mapped on a simplified phylogenetic tree. Comparison of NLR repertoires from higher plants to ancestral taxa common for plants and animals could hint at the time and mechanism which led to the assembly of NLR building blocks into a single multi-domain receptor. (25) conducted a full genome-wide comparison of NLR repertoires among 38 model organisms encompassing all the major taxa (6 eubacteria, 6 archaebacteria, 6 protists, 6 fungi, 7 plants, and 7 metazoans).Thus it is hypothesized that such a bulk control of NLR transcripts may allow a plant species to maintain large NLR repertoires without depletion of functional NLR loci (22, 23), since micro RNA-mediated transcriptional suppression of NLR transcripts could compensate for the fitness costs related to maintenance of NLRs (21, 24). The fusion events between either the NB-ARC or the NACHT domain and the LRR domain presumably occurred as indicated on the phylogenetic tree. This dataset was further enriched with the genomic and transcriptomic data available for 5,126 species of nine major early plant lineages (chlorokybales, klebsormidiales, zygnematales, coleochaetales, charales, liverworts, bryophytes, hornworts, and lycophytes).

However, host-adapted pathogens suppress this immune response by delivering effector molecules inside host cells (3, 4).

Furthermore, the demonstration of interfamily transfer of plant NLR functions from their original species to phylogenetically distant species implies evolutionary conservation of the underlying immune principle across plant taxonomy.

In this review we discuss plant NLR evolution and summarize recent insights into plant NLR-signaling mechanisms, which might constitute evolutionarily conserved NLR-mediated immune mechanisms.

The massive expansions render the NLR family one of the largest and most variable plant protein families (15, 16). (30), (d) Tarr and Alexander (31), (e) Faris et al. The number of NLR genes in flowering plants is largely variable without any clear correlation to the phylogeny, suggesting species-specific mechanisms in NLR genes expansion and/or contraction (Table 1).

This contrasts with the vertebrate NLR repertoires, typically comprising ca. Detailed genome-wide surveys, database mining, and degenerate PCR approaches for the species whose genome sequences are currently not available contribute to refine an overview of the NLR repertoires in various plant species (Table 1). This variability can be exemplified by three species in the brassicaceae family: Arabidopsis thaliana, Arabidopsis lyrata, and Brassica rapa, which have 151, 138, and 80 full-length NLRs, respectively (Table 1).

Most of the plant genomes surveyed so far have a large NLR repertoire with up to 459 genes in wine grape (Table 1). ) indicates that the presence of NLRs is not clearly resolved in given taxa due to lack of data. Expansion of NLR genes has also occurred in several metazoans such as sea urchin (Strongylocentrotus purpuratus) and sea squirt (Ciona intestinalis), which possess 206 and 203 NLRs, respectively (20, 43, 44).

Interestingly, the bryophyte Physcomitrella patens and the lycophyte Selaginella moellendorffii which represent the ancestral land plant lineages seem to have a relatively small NLR repertoire of ∼25 and ∼2 NLRs respectively, suggesting that the gene expansion has occurred mainly in flowering plants (Table 1; Figure 1). Species representative of some taxa are indicated on the right. In contrast, the genomes of fruit fly (Drosophila melanogaster) and nematode (Caenorhabditis elegans) apparently lack NLRs, suggesting that NLRs have been lost in these invertebrate species (17).

In plants and animals, nucleotide-binding domain and leucine-rich repeats (NLR)-containing proteins play pivotal roles in innate immunity.

Despite their similar biological functions and protein architecture, comparative genome-wide analyses of NLRs and genes encoding NLR-like proteins suggest that plant and animal NLRs have independently arisen in evolution.

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Analysis of the phylogeny and motif combinations of the NACHT/NB-ARC domains revealed clear differences between the NACHT and the NB-ARC domains, suggesting either an ancient divergence, or an independent origin of these two domains, which happened before the divergence of eukaryotes, eubacteria, and archaebacteria (25).

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