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The turnover of NPR1 is required for the expression of SA-responsive genes. Stabilization of NPR1, therefore, suppresses SA signaling Robert-Seilaniantz, Grant and Jones JA signaling pathways involve JAZ proteins and the SCF complex.

The bacterial toxin coronatine and the effector proteins AvrB, HopX1 and HopZ1 from P. Plant-pathogenic bacteria produce phytohormone mimics to interfere with hormone signaling pathways.

One prominent example is the phytotoxin coronatine, which is synthesized by a few pathovars of P. In addition to phytohormone mimics, plant-pathogenic bacteria deliver type III effector proteins to interfere with hormone signaling pathways.

Effectors, which interfere with JA signaling pathways, include the cysteine protease HopX1, the acetyltransferase HopZ1a and AvrB from P.

Furthermore, the cysteine protease AvrRpt2 from P. The functions of HopX1, HopZ1a, AvrB, AvrRpt2 and XopD are desribed below and summarized in Figs 5 and 6.

Modulation of JA, auxin and GA signaling pathways by type III effectors. A HopX1, HopZ1a and AvrB from P. Bioactive JA-Ile promotes the interaction between JAZ proteins and the F-box protein COI1, which is a component of the SCF complex.

The subsequent degradation of JAZ proteins leads to the release of JAZ-interacting transcription factors e. MYC2 , which activate the expression of JA-responsive genes.

The cysteine protease HopX1 directly or indirectly degrades several JAZ proteins independently of the JA receptor COI1 and thus activates the expression of JA-responsive genes.

The acetyltransferase HopZ1a acetylates JAZ proteins and leads to their proteasome-dependent degradation.

The effector protein AvrB from P. The interaction of AvrB with the RIN4-AHA1 complex promotes the interaction between JAZ proteins and COI1 and leads to the activation of JA-responsive genes.

B Auxin signaling pathways are targeted by the cysteine protease AvrRpt2 from P. ARFs subsequently activate the expression of auxin-responsive genes.

C XopD from X. GA-dependent signaling is controlled by DELLA proteins, which inactivate PIF phytochrome interacting factors transcription factors.

Binding of GA to its receptor GID1 leads to a conformational change in GID1, which subsequently binds to DELLA proteins. The formation of a GID1-DELLA complex promotes the interaction between DELLA proteins and the F-box protein SLY and thus the proteasome-dependent degradation of DELLA proteins.

This leads to the release of PIF transcription factors, which activate the expression of GA-responsive genes. XopD Xcc presumably interferes with the binding of GID1 to DELLA proteins and delays the GA-induced degradation of the DELLA protein RGA.

Notably, however, an influence of XopD Xcc on the transcription of GA-responsive genes has not yet been detected. HopX1 and HopZ1a from P.

JAZ proteins act as transcriptional repressors and interact with and inhibit transcription factors. JAZ proteins are degraded by the proteasome in the presence of JA-Ile, which is perceived by the JA receptor and F-box protein COI1 coronatine insensitive 1.

This leads to the release of JAZ-interacting transcription factors and thus to the activation of JA-responsive gene expression Robert-Seilaniantz, Grant and Jones Fig.

The effector protein HopX1 is a cysteine protease and is delivered by P. Transient coexpression studies in N. The HopX1-mediated degradation of JAZ proteins occurs independently of the JA receptor COI1 and leads to the activation of JA-responsive genes as well as to the repression of SA-induced signaling pathways.

JAZ proteins are also targeted by HopZ1a from P. HopZ1a leads to the degradation of JAZ proteins and the induction of JA-responsive genes in Arabidopsis when delivered by a coronatine-deficient mutant derivative of P.

It remains to be investigated whether the acetylation of JAZ proteins by HopZ1a facilitates their COI1-dependent degradation. The type III effector AvrB from P.

Increased concentrations of charged solutes in the guard cells result in a water uptake and elevated turgor, thus leading to stomatal opening.

RIN4 also interacts with AHA1 and promotes its activity Liu, Elmore and Coaker see below. Transient expression studies in N. The biochemical mechanism underlying the AvrB-RIN4-AHA1-mediated induction of COI1-JAZ interactions is yet unknown.

The cysteine protease AvrRpt2 from P. Members of the XopD family of nuclear-localized effector proteins from Xanthomonas spp.

Sequence comparisons revealed variations in the domain organization of XopD family members. While XopD from X. XopD Xcv deSUMOylates and thus destabilizes SlERF4 from Solanum lycopersicum Kim, Stork and Mudgett As the presence of proteasome inhibitor interferes with the XopD-induced destabilization of SlERF4, it was suggested that XopD facilitates the degradation of SlERF4 by the proteasome Kim, Stork and Mudgett SlERF4 is presumably involved in the regulation of ET biosynthesis and colocalizes with XopD to subnuclear foci Kim, Stork and Mudgett In agreement with the observed XopD-mediated destabilization of SlERF4, XopD Xcv leads to reduced ET levels in infected plant tissue and suppresses the expression of genes involved in ET production Kim, Stork and Mudgett Given that ET production is required for plant immunity, XopD Xcv likely deSUMOylates SlERF4 to suppress plant defense responses Kim, Stork and Mudgett Interference of type III effector proteins with plant gene expression.

A Domain organization of XopD proteins from Xanthomonas spp. XopD family members consist of a C-terminal cysteine protease domain and N-terminal EAR motifs.

Additionally, XopD from X. XopD Xcc was shown to interact with and stabilize DELLA proteins via the EAR motif-containing region. Furthermore, XopD Xcc interacts with and deSUMOylates the transcription factor HFR1.

XopD XccB and XopD Xcv bind to the transcription factor MYB30 via the HLH domain and suppress its transcriptional activity. XopD Xcv deSUMOylates and thus destabilizes the transcription factor ERF4.

Numbers refer to amino acid positions in XopD Xcv B Domain organization and DNA-binding specificity of the TAL effector Hax homolog of AvrBs3 in Xanthomonas 3 from X.

TAL effectors contain a C-terminal acidic activation domain AAD , two NLSs and a central protein region with repeats. The RVDs of Hax3 and the matching bases in the EBE in the promoter regions of Hax3-induced genes are indicated.

C Domain organization of HsvG from P. HsvG contains N- and C-terminal NLSs, an N-terminal HTH region and two repeats of 71 and 74 amino acids R1 and R2 , which confer transcription activation activity in yeast.

The repeats determine the specificity of plant gene activation indicated by a white arrow but are dispensable for DNA binding of HsvG, which depends on the N-terminal region.

Numbers refer to amino acid positions in HsvG. D Modification of RRS1-R by the effector protein PopP2 from R. The TIR-NB-LRR protein RRS1-R forms a dimer and binds via its WRKY domain to a DNA motif W box present in promoters of target genes of WRKY transcription factors.

PopP2 interacts with and acetylates the WRKY domain of RRS1-R and thus interferes with its DNA binding.

The additional PopP2 interaction partner RD19, which is a predicted protease, is presumably not acetylated by PopP2.

RRS1-R also interacts with the R protein RPS4, which is required for the induction of ETI and is not shown in this figure see the text for details. E The mono-ADP-RT HopU1 from P.

GRP7 also interacts with the PRRs FLS2 and EFR and with FLS2 and EFR transcripts, and was, therefore, assumed to promote PRR translation.

ADP-ribosylation of GRP7 by HopU1 reduces the ability of GRP7 to bind to RNA and might suppress FLS2 and EFR protein synthesis.

F HopD1 from P. Furthermore, HopD1 suppresses ETI responses. The mechanisms underlying the HopD1-mediated inhibition of NTL9-dependent gene expression are unknown.

DELLA proteins are negative regulators of GA response activators and colocalize with XopD Xcc to the plant nucleus.

The degradation of DELLA proteins by the proteasome is stimulated in the presence of GA, which binds to its receptor GID1 gibberellin insensitive dwarf 1 Hauvermale, Ariizumi and Steber GA promotes the interaction between GID1 and DELLA proteins, which are subsequently targeted to the F-box protein SLY and degraded by the proteasome.

The degradation of DELLA proteins leads to the activation of GA response activators, which induce the expression of GA-responsive genes Hauvermale, Ariizumi and Steber Fig.

One effective strategy employed by type III effectors to interfere with plant cellular processes is the manipulation of gene expression on the transcriptional or posttranscriptional level.

Effector proteins, which are directly imported into the nucleus and either bind to DNA or to components of the plant transcription machinery, are transcription activator-like TAL effectors from Xanthomonas spp.

Type III effectors, which target plant transcription factors and RNA-binding proteins, include XopD proteins from Xanthomonas spp.

Known mechanisms underlying type III effector-mediated modulation of plant gene expression are summarized below and in Fig. Members of the TAL effector family were mainly isolated from Xanthomonas spp.

However, related proteins are also present in R. Characteristic features of TAL effectors include a C-terminal acidic activation domain and nuclear localization signals NLSs , which are required for the import of TAL effectors into the plant nucleus Boch and Bonas DNA binding is mediated by the central region of TAL effectors, which consists of 1.

A minimum of 6. The repeats are nearly amino acid sequence identical and usually 33 to 35 amino acids long, but longer and shorter repeats have also been described Boch and Bonas Sequence-specific binding to DNA bases depends on the polymorphic amino acids at positions 12 and 13 of each repeat of the TAL effector.

The RVDs determine the binding specificity of TAL effectors to DNA. In the past years, numerous studies have focused on the analysis of the binding specificity of TAL effectors to the effector-binding elements EBEs in the promoter regions of plant target genes.

Repeat number and RVDs determine the number and nature of DNA bases, which are bound by the TAL effectors. Replacement of the natural RVDs of specific repeats by all possible RVD combinations revealed that not all artificial RVDs are functional.

In addition to the RVD composition, the length of the repeats affects the binding to DNA bases. The looping-out of repeats allows a shift of the following repeats by one nucleotide position in the EBE.

The groundbreaking discovery of the TAL-DNA-binding code has marked the beginning of a new era in genome engineering because it has led to the design of various genome editing tools e.

TAL effector nucleases , which allow sequence-specific binding of DNA-modifying enzymes by the use of TAL effector repeats as fusion partners Scharenberg, Duchateau and Smith ; Mahfouz, Piatek and Stewart The mechanisms leading to transcriptional activation of plant genes by TAL effectors are not yet understood.

It is, therefore, assumed that TAL effectors do not only bind to DNA but also associate with components of the plant transcription machinery to activate gene expression.

Among the plant genes targeted by TAL effectors are those encoding transcription factors and proteins involved in senescence, development, stress response and sugar transport Table S2, Supporting Information.

Examples are the SWEET genes from rice, which are involved in sucrose or fructose transport and are induced by TAL effectors from the systemic rice pathogen X.

TAL target genes, which contribute to virulence, are also referred to as plant susceptibility genes. Notably, however, TAL effectors can also induce the expression of plant resistance R genes and thus trigger ETI responses Boch, Bonas and Lahaye TAL effector-responsive R genes have been categorized into different groups including recessive and dominant R genes.

In this case, resistance is the result of the loss of induction of an S gene. The third group of TAL effector-responsive R genes are executor R genes, which contain EBEs in their promoter regions and are specifically activated by matching TAL effectors Zhang, Yin and White Examples are Bs3 from pepper, and Xa10 , Xa23 and Xa27 from rice Table S2.

The engineering of executor R gene promoters allows gene induction by various TAL effectors and might help to improve strategies for plant resistance and disease control Boch, Bonas and Lahaye DNA binding has also been described for the effector protein HsvG, which is an important pathogenicity factor of the gall-forming plant-pathogenic bacterium P.

HsvG is homologous to the type III effector HsvB, which is required for pathogenicity of Pa. The results of yeast one-hybrid assays suggest that HsvG acts as a transcriptional activator.

These findings suggest that the repeats and not the DNA-binding region of HsvG and HsvB determine the specificity in target gene activation.

Among the nuclear localized effectors, which presumably interfere with plant gene expression, are members of the XopD family from Xanthomonas spp.

As described above, XopD family members interfere with hormone signaling and cleave SUMO from SUMOylated proteins. It was, therefore, suggested that XopD family members suppress plant defense responses by targeting MYB Another effector protein, which modulates plant gene expression, is the YopJ family member and acetyltransferase PopP2 from R.

Notably, PopP2 does not only acetylate but also stabilizes RRS1-R. Type III effectors from plant-pathogenic bacteria do not only bind to DNA or transcription factors but can also interact with RNA-binding proteins.

One example is the mADP-RT HopU1 from P. Taken together, these findings suggest that HopU1 suppresses the GRP7-induced accumulation of FLS2 by ADP-ribosylation of GRP7.

HopD1 from P. NTL9 is a member of the NTLM1 NAC with transmembrane motif 1 family of transcription factors. This family is one of the largest families of plant transcription factors and is involved in various processes including developmental and stress-related signaling Nuruzzaman, Sharoni and Kikuchi SA is involved in various cellular processes including plant stomatal immunity, i.

Stomatal immunity is abolished in Arabidopsis ntl9 mutant plants and this phenotype is suppressed upon application of SA. Expression of NTL9-induced genes during ETI is reduced in the presence of HopD1.

As mentioned above, HopM1 from P. Taken together, these results suggest that HopM1 targets a protein to interfere with the activity of a transcriptional repressor.

The analysis of actin filaments using a GFP green fluorescent protein -fABD2 filamentous actin-binding domain 2 reporter fusion in Arabidopsis epidermis cells revealed a transient increase in actin filaments upon infection with Pseudomonas syringae pv.

A similar formation of actin filaments was observed upon treatment of plants with PAMPs and was shown to depend on FLS2, BAK1 and BIK1.

Twenty-four hours after infection with the P. No changes were induced by P. Infiltration of latrunculin B, which inhibits actin polymerization, promotes susceptibility of Arabidopsis leaves to bacterial infections and leads to an increased growth of P.

Influence of type III effectors on actin filaments and microtubules. A Infection of Arabidopsis cells with P. The infection with wild-type or T3S mutant strains leads to an increase in actin filaments 6 hours post infection.

Twenty-four hours post infection, the wild-type strain induces the formation of actin bundles and leads to a reduced number of actin filaments.

Actin filaments and bundles are indicated as yellow dashes. The plant cell wall is represented in green. The following cell organelles are shown: chloroplasts green , mitochondria beige , vacuole blue , nucleus beige , ER light brown and Golgi apparatus red.

B HopW1 leads to the disruption of actin filaments. The effector protein HopW1 binds to filamentous actin and leads to the disruption of actin filaments.

C HopG1 induces the formation of actin bundles. HopG1 binds to a mitochondrial-localized kinesin motor protein, which associates with microtubules and presumably links microtubules to actin filaments.

HopG1 induces the formation of actin bundles, presumably via its interaction with kinesin. D HopE1 leads to the dissociation of MAP65 from microtubules.

HopE1 interacts with calmodulin CaM and the microtubule-associated protein MAP65 and leads to the dissociation of MAP65 from microtubules.

No effect of HopE1 on the microtubule network was observed. E HopZ1a from P. The acetyltransferase HopZ1a binds to and acetylates tubulin and disrupts microtubules.

In addition to actin filaments, plant defense against bacterial infections also involves microtubules. Treatment of Arabidopsis plants with oryzalin, which disrupts microtubules, enhances the growth of P.

It was, therefore, suggested that effector proteins from P. The influence of type III effector proteins including HopW1, HopG1 and HopE1 from P.

HopW1 from P. Confocal microscopy using the reporter protein Lifeact-GFP, which binds to filamentous actin, showed that the delivery of HopW1 by P.

A disruption of actin filaments was also observed when hopW1 was transiently expressed in N. Thus, the delivery of HopW1 by P. The mechanisms by which HopW1 destabilizes actin remain to be elucidated.

HopG1 from P. Kinesin motor proteins are known as microtubule-associated proteins but can also interact with actin filaments and might be involved in the crosstalk and crosslinking between microtubules and actin filaments Schneider and Persson The results of coimmunprecipitation experiments suggest that HopG1 and kinesin associate with actin.

The effector HopE1 from P. No effects were observed for HopE1 derivatives with a mutated calmodulin-binding site, which is located between amino acids and of HopE1 and required for the contribution of HopE1 to in planta growth of P.

Arabidopsis map65 mutants are impaired in PTI responses and more susceptible to infections with P. The YopJ effector family members HopZ1a from P.

Fluorescence microscopy of Arabidopsis seedlings containing GFP-labeled microtubule markers revealed that delivery of HopZ1a by P. Twenty-two hours post infection, microtubules were destroyed even in the absence of HopZ1a, suggesting that microtubules are also targeted by additional effectors from P.

In agreement with the HopZ1a-mediated destruction of microtubules and the role of microtubules in vesicle trafficking, the transient expression of HopZ1a in N.

An interference with secretion of secGFP was also reported for the YopJ homolog XopJ from X. AvrBsT from X. Infection experiments revealed that the in planta growth of virulent and avirulent strains of P.

The analysis of a GFP-ACIP1 fusion revealed that ACIP1 colocalizes with microtubules. It remains to be investigated whether AvrBsT acetylates ACIP1 to interfere with plant defense responses and to alter microtubule formation.

The relevance of most of these interactions for the virulence function of AvrBsT has not yet been adressed and should be in the focus of future studies.

Type III effectors do not always act as virulence factors but can also trigger defense responses in plants, which possess corresponding R genes and can thus recognize individual effector proteins.

NLR protein activity is tightly regulated to avoid unnecessary and harmful activation of defense responses. On the posttranscriptional level, intramolecular interactions between the LRR and the NB domain of the R protein might prevent NB-mediated nucleotide exchange and thus NLR activation Bonardi and Dangl ; Takken and Goverse Activation of R protein-mediated defense responses by type III effectors.

A Detection of effector protein-triggered modifications in plant target proteins by R proteins. Type III effectors interact with and modify plant target proteins to promote bacterial virulence.

According to the guard model, plant R proteins in resistant plants detect effector-triggered modifications indicated by yellow asterisks in plant target molecules.

The activity of the NB domain is often regulated by intramolecular interactions with the LRR domain. The detection of effector-triggered modifications in plant targets leads to intramolecular rearrangements in the R proteins and thus to the activation of the NB domain indicated by a red asterisk.

According to an alternative model, plants have evolved decoy molecules, which resemble virulence targets of effectors but do not contribute to bacterial virulence.

Effector-mediated modifications in plant decoys can also lead to the activation of R protein-mediated resistance. The TIR-NB-LRR proteins RPS4 and RRS1-R form a heterodimer in the plant nucleus, and RRS1-R negatively regulates RPS4.

The WRKY domain of RRS1-R binds to DNA and presumably triggers the RRS1-R-dependent expression of plant genes. The effector proteins PopP2 from R.

It has been suggested that the WRKY domain of RRS1-R acts as an integrated decoy, which allows the elicitation of AvrRps4- and PopP2-triggered ETI responses.

The activated complex of RRS1-R and RPS4 was proposed to be a tetramer. Direct recognition of a bacterial type III effector was reported for the R protein RRS1-R from Arabidopsis which interacts with the type III effector PopP2 from R.

Most other known NLR proteins recognize their cognate effectors presumably upon detection of effector-triggered changes in plant target molecules. According to the so-called guard model, a small repertoire of R proteins can detect a wide variety of effector proteins by guarding common effector targets Van der Biezen and Jones Fig.

Alternatively, plant R proteins were proposed to detect effector-mediated changes in a non-functional effector target mimic, which acts as a decoy to trap the effector Van der Hoorn and Kamoun Fig.

Decoys might have evolved from effector targets by gene duplication and do not contribute to the pathogen's fitness. In an evolutionary point of view, the presence of decoys guarantees the persistence of NLR-mediated effector recognition, even upon changes in the virulence targets of effector proteins.

Known examples of guarded effector targets are detailed below and include i the Pto kinase, which interacts with AvrPto and AvrPtoB; ii RIN4, which is a negative regulator of plant immunity; iii the RLCK PBS1, which is cleaved by AvrPphB; and iv the pseudokinase ZED1, which mediates the recognition of HopZ1.

Effector-R protein interactions and guarded plant targets or decoys of effector proteins are also summarized in Figs 9 — One example for a putative plant decoy, which mimics an effector target, is the WRKY domain of the R protein RRS1-R, which is bound and acetylated by PopP2 from R.

It was proposed that RRS1-R acts as a sensor R protein, which binds to the effector proteins PopP2 and AvrRps4 and triggers plant defense via the executor R protein RPS4 Delga, Le Roux and Deslandes The contribution of these interactions to the virulence functions of PopP2 and AvrRps4 remains to be investigated.

The guard model was initially proposed for the recognition of the effector proteins AvrPto and AvrPtoB from P. Pto is autophosphorylated at S and required to maintain Prf in an inactive conformation.

The sensor Pto molecule interacts with and phosphorylates the E3 ubiquitin ligase AvrPtoB from P.

The Pto helper molecule transphosphorylates the sensor at amino acid residue T Given the assumed roles of AvrPto and AvrPtoB in the suppression of PTI, Pto might have evolved as a decoy to detect the activities of AvrPto and AvrPtoB inside the plant cell.

Binding of AvrPto or AvrPtoB to Pto likely disturbs the Pto-mediated negative regulation of the R protein Prf and thus leads to ETI. In many kinases, phosphorylation events within the activation segment lead to conformational changes and thus alterations in the kinase activity Nolen, Taylor and Ghosh Results of selected transient expression assays with Pto , Prf and avrPto B in N.

The promoters, which were used for transient expression or the expression of transgenes, are indicated in all cases for which this information was provided in the publications.

Pnat , native promoter of Prf ; 35S , 35S promoter; DEX , DEX-inducible promoter; transgene, integration of the gene into the genome of N.

For the better understanding of some of the results of the selected transient expression studies, conclusions provided by the authors of the indicated publications are shortly summarized.

See also the text for details. The role of the Pto kinase activity in the elicitation of ETI has been intensively studied by several research groups.

According to an alternative hypothesis, however, Pto evades the degradation by AvrPtoB by binding to two distinct binding sites in the N-terminal and central region of AvrPtoB Mathieu, Schwizer and Martin It was suggested that binding of Pto to the central region of AvrPtoB adjacent to the E3 ubiquitin ligase domain leads to the degradation of Pto.

In contrast, Pto bound to the N-terminal-binding site in AvrPtoB was reported to be stable Mathieu, Schwizer and Martin This model was mainly based on the results of yeast two-hybrid studies, which revealed that Pto does not interact with an AvrPtoB derivative containing the N-terminal Pto-binding site in fusion with the E3 ubiquitin ligase domain Mathieu, Schwizer and Martin The authors concluded from this observation that the binding of Pto adjacent to the E3 ubiquitin ligase domain of AvrPtoB leads to its AvrPtoB-dependent degradation.

Biochemical evidence for this hypothesis is still missing. Based on this finding, it was proposed that the Pto kinase activity is dispensable for the recognition of AvrPtoB and the elicitation of ETI.

This is in agreement with the model that the kinase activity of Pto is dispensable for signaling per se but required for Pto to evade degradation by AvrPtoB see above.

The double phosphorylation of Pto finally activates Prf, which triggers plant defense responses. Plant interaction partners, effector-triggered modifications of RIN4 and their effect on plant defense responses are summarized below and in Fig.

Effector-triggered modifications of RIN4 and their contributions to PTI and ETI responses. A Domain organization of RIN4 and list of known plant interaction partners of RIN4.

The PxFGxW motif is the cleavage site of the effector protein AvrRpt2 from P. Additional important amino acids are indicated see the text for details.

Known plant interaction partners of RIN4 and their predicted functions are listed. B Contribution of RIN4 to RPS2- and RPM1-triggered ETI responses.

RIN4 is also degraded by AvrPto from P. The cleavage products of RIN4 are detected by the R protein RPS2, which triggers ETI. The effector proteins AvrRpm1 and AvrB from P.

Effector-triggered phosphorylation of RIN4 presumably depends on the kinase RIPK, which interacts with RIN4 and AvrB and phosphorylates RIN4 at several amino acid residues including T Phosphorylation of RIN4 in the presence of AvrB likely induces conformational changes, which interfere with ROC1-mediated isomerization of RIN4 and lead to the activation of RPM1 see the text for details.

C Model of the role of RIN4 during PTI. The phosphorylation of RIN4 at amino acid T by AvrB and AvrRpm1 from P. RIN4 is cleaved by the effector protein AvrRpt2 from P.

At first glance, the cleavage of a PTI suppressor such as RIN4 by AvrRpt2 does not appear to be favorable for the pathogen.

However, RIN4 cleavage products were shown to be even more active PTI suppressors than the uncleaved protein Afzal, da Cunha and Mackey RIN4 is presumably not the only target of AvrRpt2 because predicted AvrRpt2 cleavage sites are also present in other plant proteins.

Interestingly, RPS2 can also be activated by the effector protein AvrRpm1 from P. As described below, the AvrRpm1-mediated phosphorylation of RIN4 is detected by the R protein RPM1.

The type III effectors AvrRpm1 and AvrB from P. The latter finding suggests that AvrB and AvrRpm1 induce the phosphorylation of RIN4 to suppress PTI responses.

RIPK is also targeted by the cysteine protease AvrPphB from P. AvrB does not only target RIPK but also interacts with MPK4, which is an additional RIN4 interaction partner see above.

The biochemical mechanisms leading to the AvrRpm1- and AvrB-induced phosphorylation of RIN4 and their impact on ETI and PTI are not yet completely understood.

Notably, however, the increased RIN4 cleavage in the presence of ROC1 S58F does not promote the RPS2-specific HR. A targeted mutagenesis approach led to the identification of P as essential residue for RPM1 activation.

It was, therefore, suggested that the activation of RPM1 does not only depend on the phosphorylation of RIN4 at T but also on P Deletion of P leads to a constitutive phosphorylation of RIN4 and thus to the activation of RPM1, suggesting that the phosphorylation of RIN4 is regulated by protein conformation.

Notably, phosphorylation of RIN4 at T leads to a reduced interaction of RIN4 with ROC1 and could thus interfere with the ROC1-mediated isomerization of P The authors of this study proposed that ROC1 induces a specific conformational change in RIN4, which interferes with the activation of RPM1 and RPS2.

Phosphorylation of RIN4 in the presence of AvrB likely leads to conformational changes that suppress the ROC1-mediated isomerization of RIN4.

Notably, the AvrB- and AvrRpm1-dependent phosphorylation of RIN4 also affects its role in PTI. Phosphorylation of S is reduced in the phosphomimic RIN4 derivative RIN4 TD , suggesting that the T phosphorylation is epistatic to S phosphorylation Fig.

The cysteine protease AvrPphB from P. It is assumed that RPS5 detects a conformational change in PBS1 and thus triggers ETI Fig. Detection of effector-triggered modifications in plant target molecules by the R proteins RPS5 and ZAR1.

A RPS5 activates ETI upon cleavage of PBS1. The CC-NB-LRR protein RPS5 interacts via the CC domain with the kinase PBS1. Cleavage of PBS1 by the effector protein and cysteine protease AvrPphB leads to the activation of RPS5.

Exchange of the AvrPphB cleavage site indicated by a yellow triangle against the recognition site of the cysteine protease AvrRpt2 indicated by an orange triangle leads to cleavage of PBS1 in the presence of AvrRpt2 and thus to the activation of RPS5.

B The R protein ZAR1 detects modifications in ZED1 and PBL2. The CC-NB-LRR protein ZAR1 interacts via the CC domain with the pseudokinase ZED1.

Acetylation of ZED1 by the effector protein HopZ1a leads to the activation of ZAR1 indicated by a red asterisk and thus to ETI responses. ZAR1 also interacts via the LRR domain with the ZED1-related pseudokinase RKS1.

The ZED1-RKS1 complex detects modifications in PBL2, which is a target of the uridylyltransferase AvrAC from X.

Uridylylation of PBL2 by AvrAC leads to the activation of ZAR1-dependent ETI responses. Cleavage of PBS1 by AvrPphB presumably leads to conformational changes in RPS5, which relieves the negative regulation of the NB by the LRR domain and thus leads to the activation of RPS5.

A recent study showed that PBS1 can be engineered to mediate recognition of bacterial effectors with protease activity other than AvrPphB. Another plant decoy is the pseudokinase ZED1 from Arabidopsis , which interacts with and is acetylated by the YopJ family member and acetyltransferase HopZ1a from P.

In addition to ZED1, ZAR1 interacts with the ZED1-related pseudokinase RKS1. Uridylylated PBL2 is recruited to the ZAR1-RKS1 complex and triggers ZAR1-dependent ETI Fig.

In order to regain his money, Zeki works during the night in the cellar of the school on a tunnel leading to the buried money. He also copies Lisi's diploma because he does not have a high school or college education, which eventually becomes known to Lisi.

She then blackmails Zeki, allowing her to get her old seventh grade class back while Zeki assumes responsibility over class 10B. Zeki is thrown out of his living situation at a strip club, and tries to sleep in Lisi's garage.

When discovered, Lisi allows Zeki to live in her basement as long as he properly teaches class 10B instead of watching films every class period.

Through unconventional methods and Lisi's soft leadership, Zeki gains the respect of the class. Although Zeki begins his time as a teacher with draconian methods including shooting students with a paintball gun and holding a student underwater, he eventually takes a softer approach and convinces the students that they do not want to become drug dealers and dependent on welfare by taking them on a field trip to visit Zeki's acquaintances who live this lifestyle.

He also becomes more involved in the school by taking over the leadership of the Drama Club, which performs a modern version of Romeo and Juliet , and helping out with the Jugend forscht group.

As a result of this increased respect, Lisi passes her practical teaching exam with class 10B. In addition, Zeki arranges an affair for Lisi's little sister Laura with her crush Danger, as well as ensuring Lisi's continued legal guardianship of Laura by pretending to be Lisi's serious boyfriend when Laura's social worker comes to visit.

Eventually Zeki finds the money in the tunnel, but the tunnel beneath the gymnasium causes the floor to break and allows Lisi to discover Zeki filling in the tunnel.

When Lisi learns about Zeki's past as a criminal, she threatens to call the police if he does not leave his job and her home at once.

Zeki, out of options, agrees to drive the get away car for a bank heist. As Zeki's class is about to take their final exam in German, they all take out their motivation photos.

A student informs Lisi that Zeki's motivation photo is in his desk. When Lisi opens the desk, she discovers that his motivation for becoming a better person is her.

A friend of Zeki's is able to convince that Zeki wants to change his ways for her. He sends Lisi a dress and an invitation to the prom and reports himself to the school principal.

The principal wants to keep him and even hands him a falsified high school diploma with a 2. She informs Zeki that the class 10B has drastically improved and if they continue to work towards their high school diplomas, the school will become the best in the city.

Their marks in German, previously 5's and 6's equivalent to F's , have now become better than 3's C's. Producer Lena Schömann praised M'Barek for his 'unbelievable discipline', as the actor had worked out five times a week with a personal trainer, losing eight kilos, months before filming began.

M'Barek had just trained his upper body for Türkisch für Anfänger. The main female role was played by Grimme Award winner Karoline Herfurth, who after Mädchen, Mädchen and its sequel was only seen rarely in comedies.

Schömann gave his reasons for Herfurth's choice, stating that she 'has a fantastic instinct for comedy and timing'. Jella Haase, who appears as Chantal, had previously been awarded the Studio Hamburg Nachwuchspreis Studio Hamburg Young Performer's Award and the Günter-Strack-Fernsehpreis Günter Strack Television Prize for best actress.

Göhre, F. Ossenbühl, M. Eichacker and J. Rochaix One of two alb3 proteins is essential for the assembly of the photosystems and for cell survival in Chlamydomonas.

Plant Cell 18 : Hu, B. Potthoff, C. Hollenberg and M. Ramezani-Rad Mdy2, a ubiquitin-like UBL -domain protein, is required for efficient mating in Saccharomyces cerevisiae.

J Cell Sci. Becht, E. Vollmeister, and M. Feldbrügge A role for RNA-binding proteins implicated in pathogenic development of Ustilago maydis.

Cell 4 : Kämper, G. Steinberg, and R. Kahmann Regulation of mating and pathogenic development in Ustilago maydis. Julius, and M.

Feldbrügge A reverse genetic approach for generating gene replacement mutants in Ustilago maydis. Ossenbühl, V. Göhre, J.

Meurer, A. Krieger-Liszkay, J. Rochaix and L. Eichacker Efficient assembly of photosystem II in Chlamydomonas reinhardtii requires Alb3. Plant Cell 16 : Müller, G.

Weinzierl, A. Feldbrügge, and R. Kahmann Mating and pathogenic development of the smut fungus Ustilago maydis are regulated by one MAP kinase cascade.

Cell 2 : Kaffarnik, P. Leibundgut, R. Feldbrügge PKA and MAPK phosphorylation of Prf1 allows promoter discrimination in Ustilago maydis. Ramezani-Rad, C.

Hollenberg, J. Lauber, H. Wedler, E. Griess, C. Wagner, K. Albermann, J. Hani, M. Piontek, U. Dahlems and G.

Gellissen The Hansenula polymorpha strain CBS Genome Sequencing and Analysis. FEMS Yeast Research 4 : Tönnis, H.

Kessler, and M. Feldbrügge Structure-function analysis of lipopeptide pheromones from the plant pathogen Ustilago maydis.

Ramezani-Rad The role of adaptor protein Stedependent regulation of the MAPKKK Ste11 in multiple signalling pathways of yeast. Genetics 43 : Bellafiore, P.

Ferris, H. Naver, V. Göhre and J. Plant Cell 14 : La Fontaine, J. Quinn, S. Nakamoto, M. Page, V. Moseley, J. Kropat and S. Merchant Copper-Dependent Iron Assimilation Pathway in the Model Photosynthetic Eukaryote Chlamydomonas reinhardtii.

Eukaryotic Cell 1 : Feldbrügge, P. Arizti, M. Sullivan, P. Zamore, J. Belasco and P. Green Comparative analysis of the plant mRNA-destabilizing element, DST, in mammalian and tobacco cells.

Plant Mol. Loubradou, A. Kahmann A homologue of the transcriptional repressor Ssn6p antagonizes cAMP signalling in Ustilago maydis. Jansen, F.

Bühring, C. Basse, P. Krüger, C. Aichinger, K. Hansson, A. Katzenberger, G. Leibbrandt, J. Torreblanca, M.

Kahmann Communication between Ustilago maydis and its host plant maize. In: IC-MPMI Congress Proceedings: Biology of Plant-Microbe Interactions Vol.

Bisseling, W. Stiekema, eds. Kahmann, G. Steinberg, C. Basse, M. Feldbrügge, and J. Kämper Ustilago maydis , the causative agent of corn smut disease.

In: Fungal Pathology J. Kronstad, ed. Kluwer Academic publishers, Dordrecht, pp. Krüger, G. Loubradou, G. Wanner, E.

Regenfelder, M. Kahmann Activation of the cAMP pathway in Ustilago maydis reduces fungal proliferation and teliospore formation in plant tumors.

Plant-Microbe Interact. Kahmann, C. Basse, and M. Feldbrügge Fungal-plant signalling in the Ustilago maydis -maize pathosystem.

Müller, C. Aichinger, M. Kahmann The MAP kinase kpp2 regulates mating and pathogenic development in Ustilago maydis.

Entian, C. Hollenberg, G. Jansen and M. Ramezani Rad et al. Ramezani Rad, G. Bühring and C. Hollenberg Ste50p is involved in regulating filamentous growth in the yeast Saccharomyces cerevisiae and associates with Ste11p.

Genetics : Sprenger, K. Hahlbrock, and B. Weisshaar PcMYB1, a novel plant protein containing a DNA-binding domain with one MYB repeat, interacts in vivo with a light-regulatory promoter unit.

Plant J. Weisshaar The transcriptional regulator CPRF1: expression analysis and gene structure. Dujon, M.

Ramezani Rad, B. Habbig, U. Hattenhorst, C. Hollenberg et al. Rad, B. Habbig, G. Jansen, U. Hattenhorst, M. Kroll and C. Hollenberg Analysis of the DNA sequence of a bp region on the left arm of yeast chromosome XV.

YEAST 13 : Xu, G. Thoma, C. Hollenberg, M. Ramezani Rad, M. Molecular Microbiology 20 : Galibert, M. Fritz, L. Kirchrath, C.

Jin, Y. Jang, M. Kim, M. Rad, L. Kirchrath, R. Seong, S. Hong, C. Hollenberg and S. Park Characterization of SFP2, a putative sulfate permease gene of Saccharomyces cerevisiae.

Rad, H. Phan, L. Kirchrath, P. Tan, T. Kirchhausen, C. Hollenberg and G. Cell Science : Sprenger, M. Dinkelbach, K. Yazaki, K. Harter, and B.

Weisshaar Functional analysis of a light-responsive plant bZIP transcriptional regulator. Plant Cell 6 : Feldmann, M.

Ramezani Rad, L. Kirchrath, G. Xu, C. Ramezani Rad, C. Kirchrath and C. YEAST 10 : Ramezani Rad and H. Katz Retention of a co-translational translocated mutant protein of carboxypeptidase Y of Saccharomyces cerevisiae in endoplasmic reticulum.

FEMS Microbiology Lett. Kawalleck, I. Somssich, M. Weisshaar Polyubiquitin gene expression and structural properties of the ubi gene in Petroselinum crispum.

Rad, G. Xu and C. Hollenberg STE50, a novel gene required for activation of conjugation at an early step of mating in Saccharomyces cerevisiae.

Oliver, Q. Agostoni-Carbone, M. Aigle, L. Alberghina, D. Alexandraki, G. Antoine, R. Anwar, J. Ballesta, P. Benit, G. Berben, E. Bergantino, N.

Biteau, P. Bolle, M. Bolotin-Fukuhara, A. Brown, A. Brown, J. Buhler, C. Carcano, G. Carignani, H. Cederberg, R. Chanet, R. Contreras, M.

Crouzet, B. Daignan-Fornier, E. Defoor, M. Delgado, J. Demolder, C. Doira, E. Dubois, B. Dujon, A. Dusterhoft, D. Erdmann, M.

Esteban, F. Fabre, C. Fairhead, G. Faye, H. Feldmann, W. Fiers, M. Francingues-Gaillard, L. Franco, L. Frontali, H. Fukuhara, L. Fuller, P. Galland, M.

Gent, D. Gigot, V. Gilliquet, N. Glansdorff, A. Goffeau, M. Grenson, P. Grisanti, L. Grivell, M. Haasemann, D. Hatat, J.

Hoenicka, J. Hegemann, C. Herbert, F. Hilger, S. Hohmann, C. Hollenberg, K. Huse, F. Iborra, K. Indje, K. Isono, C. Jacq, M.

Jacquet, C. James, J. Jauniaux, Y. Jia, A. Jimenez, A. Kelly, U. Kleinhans, P. Kreisl, G. Lanfranchi, C. Lewis, C. Lucchini, K. Lutzenkirchen, M.

Maat, L. Mallet, G. Mannhaupet, E. Martegani, A. Mathieu, C. Maurer, D. McConnell, R. McKee, F. Messenguy, H.

Mewes, F. Molemans, M. Montague, M. Muzi Falconi, L. Navas, C. Newlon, D. Noone, C. Pallier, L. Panzeri, B.

Pearson, J. Perea, P. Philippsen, A. Pierard, R. Planta, P. Plevani, B. Poetsch, F. Pohl, B. Purnelle, M. Ramezani Rad, S. Rasmussen, A.

Raynal, M. Remacha, P. Richterich, A. Roberts, F. Rodriguez, E. Sanz, I. Schaaff-Gerstenschlager, B. Scherens, B. Schweitzer, Y.

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3 Anmerkung zu “Kleine Goehre

  1. Vudogami

    Ich meine, dass Sie nicht recht sind. Ich biete es an, zu besprechen. Schreiben Sie mir in PM, wir werden umgehen.

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