Staff Publications

Staff Publications

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    'Staff publications' is the digital repository of Wageningen University & Research

    'Staff publications' contains references to publications authored by Wageningen University staff from 1976 onward.

    Publications authored by the staff of the Research Institutes are available from 1995 onwards.

    Full text documents are added when available. The database is updated daily and currently holds about 240,000 items, of which 72,000 in open access.

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    Taking stock of national climate policies to evaluate implementation of the Paris Agreement
    Roelfsema, Mark ; Soest, Heleen L. van; Harmsen, Mathijs ; Vuuren, Detlef P. van; Bertram, Christoph ; Elzen, Michel den; Höhne, Niklas ; Iacobuta, Gabriela ; Krey, Volker ; Kriegler, Elmar ; Luderer, Gunnar ; Riahi, Keywan ; Ueckerdt, Falko ; Després, Jacques ; Drouet, Laurent ; Emmerling, Johannes ; Frank, Stefan ; Fricko, Oliver ; Gidden, Matthew ; Humpenöder, Florian ; Huppmann, Daniel ; Fujimori, Shinichiro ; Fragkiadakis, Kostas ; Gi, Keii ; Keramidas, Kimon ; Köberle, Alexandre C. ; Aleluia Reis, Lara ; Rochedo, Pedro ; Schaeffer, Roberto ; Oshiro, Ken ; Vrontisi, Zoi ; Chen, Wenying ; Iyer, Gokul C. ; Edmonds, Jae ; Kannavou, Maria ; Jiang, Kejun ; Mathur, Ritu ; Safonov, George ; Vishwanathan, Saritha Sudharmma - \ 2020
    Nature Communications 11 (2020)1. - ISSN 2041-1723

    Many countries have implemented national climate policies to accomplish pledged Nationally Determined Contributions and to contribute to the temperature objectives of the Paris Agreement on climate change. In 2023, the global stocktake will assess the combined effort of countries. Here, based on a public policy database and a multi-model scenario analysis, we show that implementation of current policies leaves a median emission gap of 22.4 to 28.2 GtCO2eq by 2030 with the optimal pathways to implement the well below 2 °C and 1.5 °C Paris goals. If Nationally Determined Contributions would be fully implemented, this gap would be reduced by a third. Interestingly, the countries evaluated were found to not achieve their pledged contributions with implemented policies (implementation gap), or to have an ambition gap with optimal pathways towards well below 2 °C. This shows that all countries would need to accelerate the implementation of policies for renewable technologies, while efficiency improvements are especially important in emerging countries and fossil-fuel-dependent countries.

    Short term policies to keep the door open for Paris climate goals
    Kriegler, Elmar ; Bertram, Christoph ; Kuramochi, Takeshi ; Jakob, Michael ; Pehl, Michaja ; Stevanović, Miodrag ; Höhne, Niklas ; Luderer, Gunnar ; Minx, Jan C. ; Fekete, Hanna ; Hilaire, Jérôme ; Luna, Lisa ; Popp, Alexander ; Steckel, Jan Christoph ; Sterl, Sebastian ; Yalew, Amsalu Woldie ; Dietrich, Jan Philipp ; Edenhofer, Ottmar - \ 2018
    Environmental Research Letters 13 (2018)7. - ISSN 1748-9318
    1.5C - 2C temperature limits - Carbon pricing - Integrated assessment - Mitigation pathway - Paris Agreement - Political implementability - Regulatory policies

    Climate policy needs to account for political and social acceptance. Current national climate policy plans proposed under the Paris Agreement lead to higher emissions until 2030 than cost-effective pathways towards the Agreements' long-term temperature goals would imply. Therefore, the current plans would require highly disruptive changes, prohibitive transition speeds, and large long-term deployment of risky mitigation measures for achieving the agreement's temperature goals after 2030. Since the prospects of introducing the cost-effective policy instrument, a global comprehensive carbon price in the near-term, are negligible, we study how a strengthening of existing plans by a global roll-out of regional policies can ease the implementation challenge of reaching the Paris temperature goals. The regional policies comprise a bundle of regulatory policies in energy supply, transport, buildings, industry, and land use and moderate, regionally differentiated carbon pricing. We find that a global roll-out of these policies could reduce global CO2 emissions by an additional 10 GtCO2eq in 2030 compared to current plans. It would lead to emissions pathways close to the levels of cost-effective likely below 2C scenarios until 2030, thereby reducing implementation challenges post 2030. Even though a gradual phase-in of a portfolio of regulatory policies might be less disruptive than immediate cost-effective carbon pricing, it would perform worse in other dimensions. In particular, it leads to higher economic impacts that could become major obstacles in the long-term. Hence, such policy packages should not be viewed as alternatives to carbon pricing, but rather as complements that provide entry points to achieve the Paris climate goals.

    The Shared Socioeconomic Pathways and their energy, land use, and greenhouse gas emissions implications: An overview
    Riahi, Keywan ; Vuuren, Detlef P. Van; Kriegler, Elmar ; Edmonds, Jae ; O’neill, Brian C. ; Fujimori, Shinichiro ; Bauer, Nico ; Calvin, Katherine ; Dellink, Rob ; Fricko, Oliver ; Lutz, Wolfgang ; Popp, Alexander ; Cuaresma, Jesus Crespo ; KC, Samir ; Leimbach, Marian ; Jiang, Leiwen ; Kram, Tom ; Rao, Shilpa ; Emmerling, Johannes ; Ebi, Kristie ; Hasegawa, Tomoko ; Havlik, Petr ; Humpenöder, Florian ; Silva, Lara Aleluia Da; Smith, Steve ; Stehfest, Elke ; Bosetti, Valentina ; Eom, Jiyong ; Gernaat, David ; Masui, Toshihiko ; Rogelj, Joeri ; Strefler, Jessica ; Drouet, Laurent ; Krey, Volker ; Luderer, Gunnar ; Harmsen, Mathijs ; Takahashi, Kiyoshi ; Baumstark, Lavinia ; Doelman, Jonathan C. ; Kainuma, Mikiko ; Klimont, Zbigniew ; Marangoni, Giacomo ; Lotze-campen, Hermann ; Obersteiner, Michael ; Tabeau, Andrzej ; Tavoni, Massimo - \ 2016
    Global environmental change : human and policy dimensions 42 (2016). - ISSN 0959-3780 - p. 153 - 168.
    This paper presents the overview of the Shared Socioeconomic Pathways (SSPs) and their energy, land use, and emissions implications. The SSPs are part of a new scenario framework, established by the climate change research community in order to facilitate the integrated analysis of future climate impacts, vulnerabilities, adaptation, and mitigation. The pathways were developed over the last years as a joint community effort and describe plausible major global developments that together would lead in the future to different challenges for mitigation and adaptation to climate change. The SSPs are based on five narratives describing alternative socio-economic developments, including sustainable development, regional rivalry, inequality, fossil-fueled development, and middle-of-the-road development. The long-term demographic and economic projections of the SSPs depict a wide uncertainty range consistent with the scenario literature. A multi-model approach was used for the elaboration of the energy, land-use and the emissions trajectories of SSP-based scenarios. The baseline scenarios lead to global energy consumption of 400–1200 EJ in 2100, and feature vastly different land-use dynamics, ranging from a possible reduction in cropland area up to a massive expansion by more than 700 million hectares by 2100. The associated annual CO2 emissions of the baseline scenarios range from about 25 GtCO2 to more than 120 GtCO2 per year by 2100. With respect to mitigation, we find that associated costs strongly depend on three factors: (1) the policy assumptions, (2) the socio-economic narrative, and (3) the stringency of the target. The carbon price for reaching the target of 2.6 W/m2 that is consistent with a temperature change limit of 2 °C, differs in our analysis thus by about a factor of three across the SSP marker scenarios. Moreover, many models could not reach this target from the SSPs with high mitigation challenges. While the SSPs were designed to represent different mitigation and adaptation challenges, the resulting narratives and quantifications span a wide range of different futures broadly representative of the current literature. This allows their subsequent use and development in new assessments and research projects. Critical next steps for the community scenario process will, among others, involve regional and sectoral extensions, further elaboration of the adaptation and impacts dimension, as well as employing the SSP scenarios with the new generation of earth system models as part of the 6th climate model intercomparison project (CMIP6).
    Energy system transformations for limiting end-of-century warming to below 1.5 °C
    Rogelj, Joeri ; Luderer, Gunnar ; Pietzcker, Robert C. ; Kriegler, Elmar ; Schaeffer, Michiel ; Krey, Volker ; Riahi, Keywan - \ 2015
    Nature Climate Change 5 (2015)6. - ISSN 1758-678X - p. 519 - 527.

    Many impacts projected for a global warming level of 2 °C relative to pre-industrial levels may exceed the coping capacities of particularly vulnerable countries. Therefore, many countries advocate limiting warming to below 1.5 °C. Here we analyse integrated energy-economy-environment scenarios that keep warming to below 1.5 °C by 2100. We find that in such scenarios, energy-system transformations are in many aspects similar to 2 °C-consistent scenarios, but show a faster scale-up of mitigation action in most sectors, leading to observable differences in emission reductions in 2030 and 2050. The move from a 2 °C- to a 1.5 °C-consistent world will be achieved mainly through additional reductions of CO 2. This implies an earlier transition to net zero carbon emissions worldwide, to be achieved between 2045 and 2060. Energy efficiency and stringent early reductions are key to retain a possibility for limiting warming to below 1.5 °C by 2100. The window for achieving this goal is small and rapidly closing.

    The emissions gap report 2013 - A UNEP Synthesis Report
    Elzen, M.G.J. den; Fransen, T. ; Rogner, H.H. ; Luderer, G. ; Rogelj, J. ; Schaeffer, R. ; Neufeldt, H. ; Hoehne, N.E. ; Morgan, J. ; Olhoff, A. - \ 2013
    Nairobi, Kenya : United Nations Environment Programme - ISBN 9789280733532 - 45
    Molecular basis of plant response to microbial invasion
    Wit, P.J.G.M. de; Brandwagt, B.F. ; Burg, H.A. van den; Gabriëls, S.H.E.J. ; Hoorn, R.A.L. van der; Jong, C.F. de; Klooster, J.W. van t; Kock, M.J.D. de; Kruijt, M. ; Luderer, R. ; Munnik, T. ; Stulemeijer, I.J.E. ; Thomma, B.P.H.J. ; Vervoort, J.J.M. ; Westerink, N. ; Joosten, M.H.A.J. - \ 2004
    In: Proceedings of the 11th International Congress on Molecular Plant-Microbe Interactions, St.-Petersburg, Russia, 18-26 July 2003 / Tikhonovich, I., Lugtenberg, B., Provorov, N., St. Paul, Minnesota, USA : International Society for Molecular Plant-Microbe Interactions - ISBN 9780965462532 - p. 203 - 207.
    Gene-for-gene recognition in the Cladosporium fulvum-tomato interaction
    Wit, P.J.G.M. de; Brandwagt, B.F. ; Burg, H.A. van den; Gabriëls, S.H.E.J. ; Kock, M.J.D. de; Hoorn, R.A.L. van der; Jong, C.F. de; Klooster, J.W. van t; Kruijt, M. ; Luderer, R. ; Westerink, N. ; Joosten, M.H.A.J. - \ 2003
    Journal of Plant Pathology: rivista di patologia vegetale 85 (2003)4. - ISSN 1125-4653 - p. 275 - 275.
    Molecular basis of plant response to microbial invasion
    Wit, P.J.G.M. de; Brandwagt, B.F. ; Burg, H.A. van den; Gabriëls, S.H.E.J. ; Kock, M.J.D. de; Hoorn, R.A.L. van der; Jong, C.F. de; Klooster, J.W. van t; Kruijt, M. ; Westerink, N. ; Luderer, R. ; Vervoort, J.J.M. ; Lindhout, W.H. ; Joosten, M.H.A.J. - \ 2003
    In: Volume of Abstracts: 11-th International Congress on Molecular Plant-Microbe Interactions, St.-Petersburg, Russia, 18-26 July 2003 St.-Petersburg, Russia : All-Russia Research Institute for Agricultural Microbiology - p. 19 - 19.
    Gene-for-gene recognition in the Cladosporium fulvum-tomato interaction
    Wit, P.J.G.M. de; Brandwagt, B.F. ; Burg, H.A. van den; Gabriëls, S.H.E.J. ; Kock, M.J.D. de; Hoorn, R.A.L. van der; Jong, C.F. de; Klooster, J.W. van t; Kruijt, M. ; Luderer, R. ; Westerink, N. ; Joosten, M.H.A.J. - \ 2003
    In: Book of Abstracts of Invited Papers 8th International Congress of Plant Pathology, Christchurch, New Zealand, 2-7 February 2003 [S.l.] : S.n. - ISBN 9780864761521 - p. 77 - 77.
    The molecular basis of co-evolution between Cladosporium fulvum and tomato
    Wit, P.J.G.M. de; Brandwagt, B.F. ; Burg, H.A. van den; Cai, X. ; Hoorn, R.A.L. van der; Jong, C.F. de; Klooster, J. van 't; Kock, M.J.D. de; Kruijt, M. ; Lindhout, W.H. ; Luderer, R. ; Takken, F.L.W. ; Westerink, N. ; Vervoort, J.J.M. ; Joosten, M.H.A.J. - \ 2002
    Antonie van Leeuwenhoek: : Nederlandsch tijdschrift voor hygiëne, microbiologie en serologie 81 (2002)1-4. - ISSN 0003-6072 - p. 409 - 412.
    cladosporium - plantenziekteverwekkers - solanum - genetisch bepaalde resistentie - overgevoeligheid - cladosporium - plant pathogens - solanum - genetic resistance - hypersensitivity
    Cladosporium fulvum is a semi-biotrophic pathogen, which causes leaf mold of tomato (Lycopersicon spp.). In our laboratory this pathosystem serves as a model to study gene-for-gene interactions between plants and pathogenic fungi (Joosten & De Wit 1999). Many avirulence (Avr) genes and matching resistance (Cf) genes have been cloned and we are now beginning to understand how their products can induce an array of plant defense responses, including the classic hypersensitive response (HR). Here, we will discuss the latest results of our molecular studies on this interaction. These include the isolation of: (i) two new Avr genes, Avr2 and Avr4E, (ii) the determination of the specificity determinants within the Cf-4 and Cf-9 genes by artificial domain swaps and introduction of point mutations, (iii) the analysis of polymorphism occurring in AVR9-responsive Cf genes occurring in natural populations of L. pimpinellifolium, and finally (iv) the description of an efficient method to identify early HR-related genes.
    Manipulating signal transduction pathways in plants leading to programmed cell death
    Wit, P.J.G.M. de; Brandwagt, B.F. ; Burg, H.A. van den; Kock, M.J.D. de; Hoorn, R.A.L. van der; Jong, C.F. de; Klooster, J. van 't; Kruijt, M. ; Luderer, R. ; Westerink, N. ; Lindhout, P. ; Vervoort, J.J.M. ; Joosten, M.H.A.J. - \ 2002
    In: Book of Abstracts 9th Netherlands Biotechnology Congress, Ede, The Netherlands, 14-15 March 2002. - Wageningen, The Netherlands : Netherlands Biotechnological Society, Wageningen University, Food & Bioprocess Engineering Group, 2002 - p. 43 - 43.
    The Cladosporium fulvum-tomato interaction: elicitor proteins and their perception
    Luderer, R. - \ 2002
    Wageningen University. Promotor(en): P.J.G.M. de Wit; M.H.A.J. Joosten. - S.l. : S.n. - ISBN 9789058086273 - 103
    tomaten - plantenziekteverwekkende schimmels - pathogenesis-gerelateerde eiwitten - ziekteresistentie - virulentie - genen - solanum lycopersicum - gastheer parasiet relaties - passalora fulva - tomatoes - plant pathogenic fungi - pathogenesis-related proteins - disease resistance - virulence - genes - solanum lycopersicum - host parasite relationships - passalora fulva

    The gene-for-gene concept postulates that for every dominant gene determining resistance in the host plant, there is a corresponding dominant gene conditioning avirulence in the pathogen. The simplest way to explain the biochemical basis of this concept is direct interaction between an elicitor protein, which is encoded by an avirulence ( Avr ) gene of the pathogen, and a receptor protein, which is encoded by the matching resistance ( R ) gene of the host. Perception of the elicitor protein by the host plant subsequently leads to the activation of defence responses, often including a hypersensitive response (HR).

    The research described in this thesis is focussed on the characterisation of elicitor proteins of the fungus Cladosporium fulvum and the analysis of their perception by resistant tomato plants. A striking feature of all elicitor proteins of C. fulvum is that their mature form contains an even number of cysteine residues. These cysteine residues are thought to be involved in disulfide bridges, which are essential for proper conformation and stability of the elicitors. Mutational analysis of elicitor proteins ECP1, ECP2 and ECP5, however, revealed that the role of (the even number) of cysteine residues is more complex than anticipated, as not all cysteine residues appeared to be critical for the HR-inducing activity of the elicitor proteins (Chapter 2).

    During colonisation of the apoplastic space of tomato leaves, C. fulvum secretes elicitor proteins into the apoplast. All Cf genes, mediating resistance to particular races of C. fulvum , are predicted to encode extracytoplasmic, membrane-anchored glycoproteins that contain many leucine-rich repeats (LRRs). LRR domains are thought to be involved in protein-protein interactions. The extracellular localisation of the LRR region of the Cf proteins is consistent with a direct, extracellular perception of the corresponding elicitor proteins. To validate this hypothesis, binding studies were performed between avirulence protein AVR9 and the matching resistance protein Cf-9. Although extensive studies were performed in a multidisciplinary collaboration to prove a direct interaction between the two proteins, no specific binding between AVR9 and Cf-9 could be detected (Chapter 3). This implies that the simplest interpretation of the gene-for-gene concept, involving direct interaction of a pathogen-derived elicitor with a matching resistance gene product, does not hold for the Avr9 / Cf-9 gene pair and that at least a third component is involved in the perception of AVR9 by Cf-9.

    Also for Avr2/Cf-2 -mediated resistance a third component, Rcr3, seems to be involved. To allow dissection of the biochemical mechanism of perception of avirulence protein AVR2, we set out to clone Avr2 (Chapter 4). Avr2 cDNA was cloned based on the specific HR-inducing activity of the encoded protein in Cf2 tomato plants. Like the other Avr genes of C. fulvum , Avr2 encodes a small, secreted protein with an even number of cysteine residues. Analysis of strains of C. fulvum that are virulent on Cf2 tomato lines revealed various mutations in the Avr2 ORF that all result in the production of a truncated AVR2 protein. Interestingly, an additional modification was discovered, involving the insertion of a LINE-like element (a retrotransposable element), Cfl1, in the Avr2 ORF. Cfl1 is the first LINE-like element identified in C. fulvum and provides the first example of loss of avirulence of a plant pathogen due to insertion of a retrotransposable element in an Avr gene. Analysis of two different rcr3-mutant Cf2 tomato plants revealed that their ability to respond to AVR2 with a HR correlates with their degree of resistance to AVR2-producing strains of C. fulvum . These data support a role for Rcr3 in the perception of AVR2 by Cf-2.

    Direct perception of elicitor proteins by R proteins has been the prevailing working hypothesis to explain the biochemical basis of the gene-for-gene concept for years. The results of the research that is described in this thesis, however, do not support this hypothesis. Also for most other gene-for-gene relationships studied so far, experimental evidence appears to be more consistent with indirect perception of an AVR protein by an R protein (Chapter 5). Indirect perception implies that, beside the AVR and the R protein, at least a third component is required to induce defence responses. For several gene-for-gene relationships the nature of the putative third component is known. Although each of these components are suggested to be involved in basal defence mechanisms, their nature appears to be diverse. Hence, we argue that, although some elicitors might be directly perceived by the matching R protein, for most gene-for-gene relationships elicitor perception will turn out to be more complex.

    Molecular basis of co-evolution between Cladosporium fulvum and tomato
    Wit, P.J.G.M. de; Brandwagt, B.F. ; Burg, H.A. van den; Cai, X. ; Hoorn, R.A.L. van der; Jong, C.F. de; Klooster, J. van 't; Kock, M.J.D. de; Kruijt, M. ; Lindhout, W.H. ; Luderer, R. ; Takken, F.L.W. ; Westerink, N. ; Vervoort, J.J.M. ; Joosten, M.H.A.J. - \ 2002
    In: Biology of Plant-Microbe Interactions: Vol. 3 / Leong, S.A., Allen, C., Triplett, E.W., St. Paul, Minnesota, USA : Int. Soc. for Molecular Plant-Microbe Interactions - ISBN 9780965462525 - p. 175 - 180.
    solanum - cladosporium - passalora fulva - plantenziekteverwekkers - co-evolutie - virulentie - genen - plant pathogens - coevolution - virulence - genes
    Cladosporium fulvum overcomes Cf-2-mediated resistance by producing truncated AVR2 elicitor proteins
    Luderer, R. ; Takken, F.L.W. ; Wit, P.J.G.M. de; Joosten, M.H.A.J. - \ 2002
    Molecular Microbiology 45 (2002)3. - ISSN 0950-382X - p. 875 - 884.
    The Cf-2 gene of tomato confers resistance to strains of the biotrophic pathogenic fungus Cladosporium fulvum carrying avirulence gene Avr2. To allow dissection of the biochemical mechanism of perception of AVR2 by Cf-2, we set out to clone the Avr2 gene. Here, we report the functional cloning of Avr2 cDNA, based on the induction of a hypersensitive response (HR) by the encoded AVR2 protein in Cf2 tomato plants. Analysis of strains of C. fulvum that are virulent on Cf2 tomato lines revealed various independent frameshift mutations in the Avr2 open reading frame (ORF) and a point mutation resulting in a premature stop codon. All modifications result in the production of truncated AVR2 proteins. Interestingly, an additional modification involves the insertion of a LINE-like element, Cfl1, in the Avr2 ORF. Cfl1 is the first LINE-like element identified in C. fulvum and provides the first example of loss of avirulence of a plant pathogen caused by insertion of a retrotransposable element in an Avr gene. Rcr3 represents an additional plant protein that is specifically required for Cf-2-mediated resistance. Analysis of two different rcr3 mutant Cf2 tomato plants revealed that their ability to respond to AVR2 with a HR correlates with their degree of resistance to AVR2-producing strains of C. fulvum. These data support a role for Rcr3 in the perception of AVR2 by Cf-2.
    Functional analysis of cysteine residues of ECP elicitor proteins of the fungal tomato pathogen Cladosporium fulvum
    Luderer, R. ; Kock, M.J.D. de; Dees, R.H.L. ; Wit, P.J.G.M. de; Joosten, M.H.A.J. - \ 2002
    Molecular Plant Pathology 3 (2002)2. - ISSN 1464-6722 - p. 91 - 95.
    A striking feature of all elicitor proteins of Cladosporium fulvum that are specifically recognized by tomato is that they contain an even number of cysteine residues. These cysteine residues are thought to be involved in disulphide bridges. In this study, a mutational analysis of the cysteine residues of ECP1, ECP2 and ECP5 was performed, to examine their role in stability and hypersensitive response-inducing activity of the proteins. We show that not all cysteine residues of the ECPs are critical for the hypersensitive response-inducing activity of the proteins, and we propose that the role of cysteine residues in the ECPs is more complex than anticipated.
    The role of secreted proteins of the pathogenic fungus Cladosporium fulvum in (a)virulence on its host, tomato
    Joosten, M.H.A.J. ; Brandwagt, B.F. ; Hoorn, R.A.L. van der; Jong, C.F. de; Klooster, J.W. van 't; Kock, M.J.D. de; Kruijt, M. ; Luderer, R. ; Takken, F.L.W. ; Westerink, N. ; Wit, P.J.G.M. de - \ 2001
    Phytopathology 91 (2001)6. - ISSN 0031-949X - p. S161 - S161.
    Cladosporium fulvum is a biotrophic fungal pathogen, which secretes various low molecular weight proteins into the apoplast of tomato leaves. Some of these proteins are secreted by all strains of the fungus and are essential virulence factors called extra cellular proteins or ECPs, whereas others are race-specific avirulence factors (AVRs) that induce a hypersensitive response in tomato containing the matching resistance gene. All Ecp and Avr genes cloned to date encode proteins with a typical signal sequence for extracellular targeting and contain an even number of cysteine residues (4, 6 or 8). My presentation will focus on how Ecp and Avr genes have been isolated, how we think the encoded proteins are perceived by resistant plants and what kind of responses are induced in such plants
    Avirulence proteins of plant pathogens: determinants of victory and defeat
    Luderer, R. ; Joosten, M.H.A.J. - \ 2001
    Molecular Plant Pathology 2 (2001)6. - ISSN 1464-6722 - p. 355 - 364.
    The simplest way to explain the biochemical basis of the gene-for-gene concept is by direct interaction between a pathogen-derived avirulence (Avr) gene product and a receptor protein, which is encoded by the matching resistance (R) gene of the host plant. The number of R genes for which the matching Avr gene has been cloned is increasing. The number of host-pathogen relationships, however, for which a direct interaction between R and Avr gene products could be proven is still very limited. This observation suggests that in various host-pathogen relationships no physical interaction between R and Avr proteins occurs, and that perception of AVR proteins by their matching R gene products is indirect. Indirect perception implies that at least a third component is required. The 'Guard hypothesis' proposes that this third component could be the virulence target of an AVR protein. Binding of the AVR protein to its virulence target is perceived by the matching R protein, which is 'guarding' the virulence target. An intriguing aspect of the 'Guard hypothesis' is that the Avr gene product causes avirulence of the pathogen through interaction with its virulence target in the plant. This would mean that, although AVR proteins are generally thought to be bifunctional (avirulence as well as virulence factors), this dual function might be based on a single biochemical event. This review focuses on the way AVR proteins are perceived by their matching R gene products. The various components that determine the outcome of the interaction will be discussed, with an emphasis on the dual function of AVR proteins.
    Molecular basis of co-evolution between Cladosporium fulvum and tomato
    Wit, P.J.G.M. de; Brandwagt, B. ; Hoorn, R. Van der; Jong, C. de; Klooster, J. Van 't; Kock, M. De; Kruijt, M. ; Luderer, R. ; Westerink, N. ; Joosten, M.H.A.J. - \ 2001
    In: Book of Abstracts: 10th International Congress on Molecular Plant-Microbe Interactions, Madison, WI, USA, 10-14 July 2001. - [S.l.] : [s.n.], 2001 - p. 19 - 19.
    Cloning and characterisation of avirulence gene Avr2 of Cladosporium fulvum
    Luderer, R. ; Takken, F.L.W. ; Wit, P.J.G.M. de; Joosten, M.H.A.J. - \ 2001
    In: Book of Abstracts XXI Fungal Genetics Conference, Asilomar, USA, 13-18 March 2001. - Asilomar, USA : [s.n.], 2001 - p. 95 - 95.
    No evidence for binding between resistance gene product Cf-9 of tomato and avirulence gene product AVR9 of Cladosporium fulvum
    Luderer, R. ; Rivas, S. ; Nurnberger, T. ; Mattei, B. ; Hooven, H.W. Van den; Hoorn, R.A.L. Van der; Romeis, T. ; Wehrfritz, J.M. ; Blume, B. ; Nennstiel, D. ; Zuidema, D. ; Vervoort, J.J.M. ; Lorenzo, G. De; Jones, J.D.G. ; Wit, P.J.G.M. De; Joosten, M.H.A.J. - \ 2001
    Molecular Plant-Microbe Interactions 14 (2001)7. - ISSN 0894-0282 - p. 867 - 876.
    The gene-for-gene model postulates that for every gene determining resistance in the host plant, there is a corresponding gene conditioning avirulence in the pathogen. On the basis of this relationship, products of resistance (R) genes and matching avirulence (Avr) genes are predicted to interact. Here, we report on binding studies between the R gene product Cf-9 of tomato and the Avr gene product AVR9 of the pathogenic fungus Cladosporium fulvum. Because a high-affinity binding site (HABS) for AVR9 is present in tomato lines, with or without the Cf-9 resistance gene, as well as in other solanaceous plants, the Cf-9 protein was produced in COS and insect cells in order to perform binding studies in the absence of the HABS. Binding studies with radio-labeled AVR9 were performed with Cf-9-producing COS and insect cells and with membrane preparations of such cells. Furthermore, the Cf-9 gene was introduced in tobacco, which is known to be able to produce a functional Cf-9 protein. Binding of AVR9 to Cf-9 protein produced in tobacco was studied employing surface plasmon resonance and surface-enhanced laser desorption and ionization. Specific binding between Cf-9 and AVR9 was not detected with any of the procedures. The implications of this observation are discussed.
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