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Papers by Marcel Proveniers

Research paper thumbnail of Repression of lateral organ boundary genes by PENNYWISE and POUND-FOOLISH is essential for meristem maintenance and flowering in Arabidopsis thaliana1

Plant physiology, Jan 28, 2015

In the model plant Arabidopsis (Arabidopsis thaliana), endogenous and environmental signals actin... more In the model plant Arabidopsis (Arabidopsis thaliana), endogenous and environmental signals acting on the shoot apical meristem cause acquisition of inflorescence meristem fate. This results in new patterns of aerial development, seen as the transition from making leaves to the production of flowers separated by elongated internodes. Specification of floral meristem identity requires the activities of two related BEL1-like homeodomain proteins, PENNYWISE (PNY) and POUND-FOOLISH (PNF). Loss-of-function of pny pnf mutations impair stem cell maintenance, and the ability to initiate internodes and flowers during shoot development. We show here that misexpression of lateral organ boundary genes BLADE-ON-PETIOLE1/2 (BOP1/2) and KNOTTED-like from ARABIDOPSIS THALIANA6 (KNAT6) together with ARABIDOPSIS THALIANA HOMEOBOX GENE1 (ATH1) in pny pnf apices is responsible for these defects. Inactivation of genes in this module fully rescues pny pnf defects. We show that BOP1 directly activates ATH...

Research paper thumbnail of Ethylene-induced hyponastic growth in Arabidopsis thaliana is controlled by ERECTA

Plant Journal, 2010

Plants can respond quickly and profoundly to detrimental changes in their environment. For exampl... more Plants can respond quickly and profoundly to detrimental changes in their environment. For example, Arabidopsis thaliana can induce an upward leaf movement response through differential petiole growth (hyponastic growth) to outgrow complete submergence. This response is induced by accumulation of the phytohormone ethylene in the plant. Currently, only limited information is available on how this response is molecularly controlled. In

Research paper thumbnail of Organisatie

Research paper thumbnail of Sugars speed up the circle of life

eLife, 2013

By regulating the expression of key microRNA molecules, sugar levels in leaves control the transi... more By regulating the expression of key microRNA molecules, sugar levels in leaves control the transition from the juvenile to the adult form in plants.

Research paper thumbnail of LIGHT-REGULATED TRANSCRIPTION FACTOR GENES FROM ARABIDOPSIS-THALIANA

Research paper thumbnail of The Arabidopsis homeobox gene ATH1 and floral transition

Research paper thumbnail of Plant gene constructs and their use

Research paper thumbnail of High temperature acclimation through PIF4 signaling

Trends in Plant Science, 2013

Ambient temperature has direct consequences for plant functioning. Many plant species are able to... more Ambient temperature has direct consequences for plant functioning. Many plant species are able to adjust reproductive timing and development to optimize fitness to changes in ambient temperatures. Understanding the molecular networks of how plants cope with high temperatures is essential to counteract the effects of global warming and to secure future crop productivity. Several recent papers reported that Arabidopsis thaliana responses to changing light conditions and high temperature, and their underlying signaling mechanisms are highly similar and involve the basic helix-loop-helix (bHLH) transcription factor PHYTOCHROME INTERACTING FACTOR 4 (PIF4). In this opinion article we discuss the mechanisms of PIF4-mediated acclimation to increased ambient temperature with focus on timing of flowering and morphological acclimation.

Research paper thumbnail of The many functions of ERECTA

Trends in Plant Science, 2009

The Arabidopsis thaliana accession Landsberg erecta contains an induced mutation in the leucine-r... more The Arabidopsis thaliana accession Landsberg erecta contains an induced mutation in the leucine-rich repeat receptor-like Ser/Thr kinase gene ERECTA. Landsberg erecta is commonly used as a genetic background in mutant screens and in natural variation studies. Therefore, the erecta mutation is present in many loss-of-function mutants and recombinant inbred lines. Information on how the absence of functional ERECTA affects the interpretation of obtained phenotypic results is scattered. In this report we inventoried ERECTA functions and highlight ERECTA as a pleiotropic regulator of developmental and physiological processes, as well as a modulator of responses to environmental stimuli.

Research paper thumbnail of Ethylene-induced hyponastic growth in Arabidopsis thaliana is controlled by ERECTA

The Plant Journal, 2010

Plants can respond quickly and profoundly to detrimental changes in their environment. For exampl... more Plants can respond quickly and profoundly to detrimental changes in their environment. For example, Arabidopsis thaliana can induce an upward leaf movement response through differential petiole growth (hyponastic growth) to outgrow complete submergence. This response is induced by accumulation of the phytohormone ethylene in the plant. Currently, only limited information is available on how this response is molecularly controlled. In this study, we utilized quantitative trait loci (QTL) analysis of natural genetic variation among Arabidopsis accessions to isolate novel factors controlling constitutive petiole angles and ethylene-induced hyponastic growth. Analysis of mutants in various backgrounds and complementation analysis of naturally occurring mutant accessions provided evidence that the leucin-rich repeat receptor-like Ser/Thr kinase gene, ERECTA, controls ethylene-induced hyponastic growth. Moreover, ERECTA controls leaf positioning in the absence of ethylene treatment. Our data demonstrate that this is not due to altered ethylene production or sensitivity.

Research paper thumbnail of The Arabidopsis TALE homeobox gene ATH1 controls floral competency through positive regulation of FLC

The Plant Journal, 2007

Floral induction is controlled by a plethora of genes acting in different pathways that either re... more Floral induction is controlled by a plethora of genes acting in different pathways that either repress or promote floral transition at the shoot apical meristem (SAM). During vegetative development high levels of floral repressors maintain the Arabidopsis SAM as incompetent to respond to promoting factors. Among these repressors, FLOWERING LOCUS C (FLC) is the most prominent. The processes underlying downregulation of FLC in response to environmental and developmental signals have been elucidated in considerable detail. However, the basal induction of FLC and its upregulation by FRIGIDA (FRI) are still poorly understood. Here we report the functional characterization of the ARABIDOPSIS THALIANA HOMEOBOX 1 (ATH1) gene. A function of ATH1 in floral repression is suggested by a gradual downregulation of ATH1 in the SAM prior to floral transition. Further evidence for such a function of ATH1 is provided by the vernalization-sensitive late flowering of plants that constitutively express ATH1. Analysis of lines that differ in FRI and/or FLC allele strength show that this late flowering is caused by upregulation of FLC as a result of synergism between ATH1 overexpression and FRI. Lack of ATH1, however, results in attenuated FLC levels independently of FRI, suggesting that ATH1 acts as a general activator of FLC expression. This is further corroborated by a reduction of FLC-mediated late flowering in fca-1 and fve-1 autonomous pathway backgrounds when combined with ath1. Since other floral repressors of the FLC clade are not significantly affected by ATH1, we conclude that ATH1 controls floral competency as a specific activator of FLC expression.

Research paper thumbnail of Shoot apical meristem function in Arabidopsis requires the combined activities of three BEL1-like homeodomain proteins

The Plant Journal, 2009

In plants, most of the above-ground body is formed post-embryonically by the continuous organogen... more In plants, most of the above-ground body is formed post-embryonically by the continuous organogenic potential of the shoot apical meristem (SAM). Proper function of the SAM requires maintenance of a delicate balance between the depletion of stem cell daughters into developing primordia and proliferation of the central stem cell population. Here we show that initiation and maintenance of the Arabidopsis SAM, including that of floral meristems, requires the combinatorial action of three members of the BELL-family of TALE homeodomain proteins, ARABIDOPSIS THALIANA HOMEOBOX 1 (ATH1), PENNYWISE (PNY) and POUND-FOOLISH (PNF). All three proteins interact with the KNOX TALE homeodomain protein STM, and combined lesions in ATH1, PNY and PNF result in a phenocopy of stm mutations. Therefore, we propose that ath1 pny pnf meristem defects result from loss of combinatorial BELL-STM control. Further, we demonstrate that heterodimerization-controlled cellular localization of BELL and KNOX proteins involves a CRM1/exportin-1mediated nuclear exclusion mechanism that is probably generic to control the activity of BELL and KNOX combinations. We conclude that in animals and plants corresponding mechanisms regulate the activity of TALE homeodomain proteins through controlled nuclear-cytosolic distribution of these proteins.

Research paper thumbnail of Arabidopsis Class I KNOTTED-Like Homeobox Proteins Act Downstream in the IDA-HAE/HSL2 Floral Abscission Signaling Pathway

The Plant Cell, 2011

Floral organ abscission in Arabidopsis thaliana is regulated by the putative ligand-receptor syst... more Floral organ abscission in Arabidopsis thaliana is regulated by the putative ligand-receptor system comprising the signaling peptide INFLORESCENCE DEFICIENT IN ABSCISSION (IDA) and the two receptor-like kinases HAESA and HAESA-LIKE2. The IDA signaling pathway presumably activates a MITOGEN-ACTIVATED PROTEIN KINASE (MAPK) cascade to induce separation between abscission zone (AZ) cells. Misexpression of IDA effectuates precocious floral abscission and ectopic cell separation in latent AZ cell regions, which suggests that negative regulators are in place to prevent unrestricted and untimely AZ cell separation. Through a screen for mutations that restore floral organ abscission in ida mutants, we identified three new mutant alleles of the KNOTTED-LIKE HOMEOBOX gene BREVIPEDICELLUS (BP)/KNOTTED-LIKE FROM ARABIDOPSIS THALIANA1 (KNAT1). Here, we show that bp mutants, in addition to shedding their floral organs prematurely, have phenotypic commonalities with plants misexpressing IDA, such as enlarged AZ cells. We propose that BP/KNAT1 inhibits floral organ cell separation by restricting AZ cell size and number and put forward a model whereby IDA signaling suppresses BP/KNAT1, which in turn allows KNAT2 and KNAT6 to induce floral organ abscission.

Research paper thumbnail of Isolation and characterization of mRNAs accumulated during in-vitro flower bud formation

Planta, 1994

The development of vegetative and generative buds on thin-layer explants of tobacco (Nicotiana ta... more The development of vegetative and generative buds on thin-layer explants of tobacco (Nicotiana tabacum L. cv. Samsun) has been studied at the level of translatable mRNA to detect changes in the mRNA population during bud initiation and differentiation, and several quantitative differences were found. By differential screening of a cDNA library obtained from flower-budregenerating explants we have isolated a group of six cDNA clones representing genes that are preferentially expressed during in-vitro flower bud formation. Nucleotide sequence analysis of one of these cDNAs, pAP8, showed that the most likely open reading frame has some typical characteristics of, and homology with, extensinlike genes. Northern blot analysis and in-situ hybridization suggest a specific role for these extensin-like genes in flower bud initiation on tobacco pedicel explants.

Research paper thumbnail of ERECTA controls low light intensity-induced differential petiole growth independent of Phytochrome B and Cryptochrome 2 action in A rabidopsis thaliana

Plant Signaling & Behavior, 2010

Research paper thumbnail of MYB72 Is Required in Early Signaling Steps of Rhizobacteria-Induced Systemic Resistance in Arabidopsis

PLANT PHYSIOLOGY, 2008

Pseudomonas fluorescens WCS417r bacteria triggers a jasmonate/ethylene-dependent induced systemic... more Pseudomonas fluorescens WCS417r bacteria triggers a jasmonate/ethylene-dependent induced systemic resistance (ISR) that is effective against a broad range of pathogens. Microarray analysis revealed that the R2R3-MYB-like transcription factor gene MYB72 is specifically activated in the roots upon colonization by WCS417r. Here we show that T-DNA knockout mutants myb72-1 and myb72-2 are incapable of mounting ISR against the pathogens Pseudomonas syringae pv. tomato, Hyaloperonospora parasitica, Alternaria brassicicola and Botrytis cinerea, indicating that MYB72 is essential to establish broad-spectrum ISR. Overexpression of MYB72 did not result in enhanced resistance against any of the pathogens tested, demonstrating that MYB72 is not sufficient for the expression of ISR. Yeast two-hybrid analysis revealed that MYB72 physically interacts in vitro with the ETHYLENE INSENSITIVE3 (EIN3)-LIKE3 transcription factor EIL3, linking MYB72 function to the ethylene response pathway. However, WCS417r activated MYB72 in ISR-deficient, ethylene-insensitive ein2-1 plants. Moreover, exogenous application of the ethylene precursor 1-aminocyclopropane-1-carboxylate (ACC) induced wild-type levels of resistance in myb72-1, suggesting that MYB72 acts upstream of ethylene in the ISR pathway. Collectively, this study identified the transcriptional regulator MYB72 as a novel ISR signaling component that is required in the roots during early signaling steps of rhizobacteriamediated ISR. www.plant.org on November 2, 2015 -Published by www.plantphysiol.org Downloaded from

Research paper thumbnail of Late heading of perennial ryegrass caused by introducing an Arabidopsis homeobox gene

Plant Breeding, 2004

Perennial ryegrass (Lolium perenne L.) is the most important temperate forage grass species. Unfo... more Perennial ryegrass (Lolium perenne L.) is the most important temperate forage grass species. Unfortunately, the nutritional value of perennial ryegrass declines as maturity progresses, mainly because of a high concentration of poorly digestible compounds in inflorescences. Therefore, the development of forage-type ryegrass varieties with extended vegetative growth is of interest for agriculture. To delay floral transition in perennial ryegrass the Arabidopsis ATH1 gene driven by the maize ubiquitin promoter, the rice actin promoter or the rice OSH1 promoter, respectively was introduced. In ATH1-expressing plants heading was delayed, and in a number of cases the plants never flowered at all. Such non-or late-heading was accompanied by the outgrowth of normally quiescent lateral meristems into extra leaves, resulting in a leafy growth habit. When eventually heading, these plants generally produced a reduced number of inflorescences. These observations suggest that ATH1-mediated delay of heading may be useful to improve fodder quality of perennial ryegrass.

Research paper thumbnail of Repression of lateral organ boundary genes by PENNYWISE and POUND-FOOLISH is essential for meristem maintenance and flowering in Arabidopsis thaliana1

Plant physiology, Jan 28, 2015

In the model plant Arabidopsis (Arabidopsis thaliana), endogenous and environmental signals actin... more In the model plant Arabidopsis (Arabidopsis thaliana), endogenous and environmental signals acting on the shoot apical meristem cause acquisition of inflorescence meristem fate. This results in new patterns of aerial development, seen as the transition from making leaves to the production of flowers separated by elongated internodes. Specification of floral meristem identity requires the activities of two related BEL1-like homeodomain proteins, PENNYWISE (PNY) and POUND-FOOLISH (PNF). Loss-of-function of pny pnf mutations impair stem cell maintenance, and the ability to initiate internodes and flowers during shoot development. We show here that misexpression of lateral organ boundary genes BLADE-ON-PETIOLE1/2 (BOP1/2) and KNOTTED-like from ARABIDOPSIS THALIANA6 (KNAT6) together with ARABIDOPSIS THALIANA HOMEOBOX GENE1 (ATH1) in pny pnf apices is responsible for these defects. Inactivation of genes in this module fully rescues pny pnf defects. We show that BOP1 directly activates ATH...

Research paper thumbnail of Ethylene-induced hyponastic growth in Arabidopsis thaliana is controlled by ERECTA

Plant Journal, 2010

Plants can respond quickly and profoundly to detrimental changes in their environment. For exampl... more Plants can respond quickly and profoundly to detrimental changes in their environment. For example, Arabidopsis thaliana can induce an upward leaf movement response through differential petiole growth (hyponastic growth) to outgrow complete submergence. This response is induced by accumulation of the phytohormone ethylene in the plant. Currently, only limited information is available on how this response is molecularly controlled. In

Research paper thumbnail of Organisatie

Research paper thumbnail of Sugars speed up the circle of life

eLife, 2013

By regulating the expression of key microRNA molecules, sugar levels in leaves control the transi... more By regulating the expression of key microRNA molecules, sugar levels in leaves control the transition from the juvenile to the adult form in plants.

Research paper thumbnail of LIGHT-REGULATED TRANSCRIPTION FACTOR GENES FROM ARABIDOPSIS-THALIANA

Research paper thumbnail of The Arabidopsis homeobox gene ATH1 and floral transition

Research paper thumbnail of Plant gene constructs and their use

Research paper thumbnail of High temperature acclimation through PIF4 signaling

Trends in Plant Science, 2013

Ambient temperature has direct consequences for plant functioning. Many plant species are able to... more Ambient temperature has direct consequences for plant functioning. Many plant species are able to adjust reproductive timing and development to optimize fitness to changes in ambient temperatures. Understanding the molecular networks of how plants cope with high temperatures is essential to counteract the effects of global warming and to secure future crop productivity. Several recent papers reported that Arabidopsis thaliana responses to changing light conditions and high temperature, and their underlying signaling mechanisms are highly similar and involve the basic helix-loop-helix (bHLH) transcription factor PHYTOCHROME INTERACTING FACTOR 4 (PIF4). In this opinion article we discuss the mechanisms of PIF4-mediated acclimation to increased ambient temperature with focus on timing of flowering and morphological acclimation.

Research paper thumbnail of The many functions of ERECTA

Trends in Plant Science, 2009

The Arabidopsis thaliana accession Landsberg erecta contains an induced mutation in the leucine-r... more The Arabidopsis thaliana accession Landsberg erecta contains an induced mutation in the leucine-rich repeat receptor-like Ser/Thr kinase gene ERECTA. Landsberg erecta is commonly used as a genetic background in mutant screens and in natural variation studies. Therefore, the erecta mutation is present in many loss-of-function mutants and recombinant inbred lines. Information on how the absence of functional ERECTA affects the interpretation of obtained phenotypic results is scattered. In this report we inventoried ERECTA functions and highlight ERECTA as a pleiotropic regulator of developmental and physiological processes, as well as a modulator of responses to environmental stimuli.

Research paper thumbnail of Ethylene-induced hyponastic growth in Arabidopsis thaliana is controlled by ERECTA

The Plant Journal, 2010

Plants can respond quickly and profoundly to detrimental changes in their environment. For exampl... more Plants can respond quickly and profoundly to detrimental changes in their environment. For example, Arabidopsis thaliana can induce an upward leaf movement response through differential petiole growth (hyponastic growth) to outgrow complete submergence. This response is induced by accumulation of the phytohormone ethylene in the plant. Currently, only limited information is available on how this response is molecularly controlled. In this study, we utilized quantitative trait loci (QTL) analysis of natural genetic variation among Arabidopsis accessions to isolate novel factors controlling constitutive petiole angles and ethylene-induced hyponastic growth. Analysis of mutants in various backgrounds and complementation analysis of naturally occurring mutant accessions provided evidence that the leucin-rich repeat receptor-like Ser/Thr kinase gene, ERECTA, controls ethylene-induced hyponastic growth. Moreover, ERECTA controls leaf positioning in the absence of ethylene treatment. Our data demonstrate that this is not due to altered ethylene production or sensitivity.

Research paper thumbnail of The Arabidopsis TALE homeobox gene ATH1 controls floral competency through positive regulation of FLC

The Plant Journal, 2007

Floral induction is controlled by a plethora of genes acting in different pathways that either re... more Floral induction is controlled by a plethora of genes acting in different pathways that either repress or promote floral transition at the shoot apical meristem (SAM). During vegetative development high levels of floral repressors maintain the Arabidopsis SAM as incompetent to respond to promoting factors. Among these repressors, FLOWERING LOCUS C (FLC) is the most prominent. The processes underlying downregulation of FLC in response to environmental and developmental signals have been elucidated in considerable detail. However, the basal induction of FLC and its upregulation by FRIGIDA (FRI) are still poorly understood. Here we report the functional characterization of the ARABIDOPSIS THALIANA HOMEOBOX 1 (ATH1) gene. A function of ATH1 in floral repression is suggested by a gradual downregulation of ATH1 in the SAM prior to floral transition. Further evidence for such a function of ATH1 is provided by the vernalization-sensitive late flowering of plants that constitutively express ATH1. Analysis of lines that differ in FRI and/or FLC allele strength show that this late flowering is caused by upregulation of FLC as a result of synergism between ATH1 overexpression and FRI. Lack of ATH1, however, results in attenuated FLC levels independently of FRI, suggesting that ATH1 acts as a general activator of FLC expression. This is further corroborated by a reduction of FLC-mediated late flowering in fca-1 and fve-1 autonomous pathway backgrounds when combined with ath1. Since other floral repressors of the FLC clade are not significantly affected by ATH1, we conclude that ATH1 controls floral competency as a specific activator of FLC expression.

Research paper thumbnail of Shoot apical meristem function in Arabidopsis requires the combined activities of three BEL1-like homeodomain proteins

The Plant Journal, 2009

In plants, most of the above-ground body is formed post-embryonically by the continuous organogen... more In plants, most of the above-ground body is formed post-embryonically by the continuous organogenic potential of the shoot apical meristem (SAM). Proper function of the SAM requires maintenance of a delicate balance between the depletion of stem cell daughters into developing primordia and proliferation of the central stem cell population. Here we show that initiation and maintenance of the Arabidopsis SAM, including that of floral meristems, requires the combinatorial action of three members of the BELL-family of TALE homeodomain proteins, ARABIDOPSIS THALIANA HOMEOBOX 1 (ATH1), PENNYWISE (PNY) and POUND-FOOLISH (PNF). All three proteins interact with the KNOX TALE homeodomain protein STM, and combined lesions in ATH1, PNY and PNF result in a phenocopy of stm mutations. Therefore, we propose that ath1 pny pnf meristem defects result from loss of combinatorial BELL-STM control. Further, we demonstrate that heterodimerization-controlled cellular localization of BELL and KNOX proteins involves a CRM1/exportin-1mediated nuclear exclusion mechanism that is probably generic to control the activity of BELL and KNOX combinations. We conclude that in animals and plants corresponding mechanisms regulate the activity of TALE homeodomain proteins through controlled nuclear-cytosolic distribution of these proteins.

Research paper thumbnail of Arabidopsis Class I KNOTTED-Like Homeobox Proteins Act Downstream in the IDA-HAE/HSL2 Floral Abscission Signaling Pathway

The Plant Cell, 2011

Floral organ abscission in Arabidopsis thaliana is regulated by the putative ligand-receptor syst... more Floral organ abscission in Arabidopsis thaliana is regulated by the putative ligand-receptor system comprising the signaling peptide INFLORESCENCE DEFICIENT IN ABSCISSION (IDA) and the two receptor-like kinases HAESA and HAESA-LIKE2. The IDA signaling pathway presumably activates a MITOGEN-ACTIVATED PROTEIN KINASE (MAPK) cascade to induce separation between abscission zone (AZ) cells. Misexpression of IDA effectuates precocious floral abscission and ectopic cell separation in latent AZ cell regions, which suggests that negative regulators are in place to prevent unrestricted and untimely AZ cell separation. Through a screen for mutations that restore floral organ abscission in ida mutants, we identified three new mutant alleles of the KNOTTED-LIKE HOMEOBOX gene BREVIPEDICELLUS (BP)/KNOTTED-LIKE FROM ARABIDOPSIS THALIANA1 (KNAT1). Here, we show that bp mutants, in addition to shedding their floral organs prematurely, have phenotypic commonalities with plants misexpressing IDA, such as enlarged AZ cells. We propose that BP/KNAT1 inhibits floral organ cell separation by restricting AZ cell size and number and put forward a model whereby IDA signaling suppresses BP/KNAT1, which in turn allows KNAT2 and KNAT6 to induce floral organ abscission.

Research paper thumbnail of Isolation and characterization of mRNAs accumulated during in-vitro flower bud formation

Planta, 1994

The development of vegetative and generative buds on thin-layer explants of tobacco (Nicotiana ta... more The development of vegetative and generative buds on thin-layer explants of tobacco (Nicotiana tabacum L. cv. Samsun) has been studied at the level of translatable mRNA to detect changes in the mRNA population during bud initiation and differentiation, and several quantitative differences were found. By differential screening of a cDNA library obtained from flower-budregenerating explants we have isolated a group of six cDNA clones representing genes that are preferentially expressed during in-vitro flower bud formation. Nucleotide sequence analysis of one of these cDNAs, pAP8, showed that the most likely open reading frame has some typical characteristics of, and homology with, extensinlike genes. Northern blot analysis and in-situ hybridization suggest a specific role for these extensin-like genes in flower bud initiation on tobacco pedicel explants.

Research paper thumbnail of ERECTA controls low light intensity-induced differential petiole growth independent of Phytochrome B and Cryptochrome 2 action in A rabidopsis thaliana

Plant Signaling & Behavior, 2010

Research paper thumbnail of MYB72 Is Required in Early Signaling Steps of Rhizobacteria-Induced Systemic Resistance in Arabidopsis

PLANT PHYSIOLOGY, 2008

Pseudomonas fluorescens WCS417r bacteria triggers a jasmonate/ethylene-dependent induced systemic... more Pseudomonas fluorescens WCS417r bacteria triggers a jasmonate/ethylene-dependent induced systemic resistance (ISR) that is effective against a broad range of pathogens. Microarray analysis revealed that the R2R3-MYB-like transcription factor gene MYB72 is specifically activated in the roots upon colonization by WCS417r. Here we show that T-DNA knockout mutants myb72-1 and myb72-2 are incapable of mounting ISR against the pathogens Pseudomonas syringae pv. tomato, Hyaloperonospora parasitica, Alternaria brassicicola and Botrytis cinerea, indicating that MYB72 is essential to establish broad-spectrum ISR. Overexpression of MYB72 did not result in enhanced resistance against any of the pathogens tested, demonstrating that MYB72 is not sufficient for the expression of ISR. Yeast two-hybrid analysis revealed that MYB72 physically interacts in vitro with the ETHYLENE INSENSITIVE3 (EIN3)-LIKE3 transcription factor EIL3, linking MYB72 function to the ethylene response pathway. However, WCS417r activated MYB72 in ISR-deficient, ethylene-insensitive ein2-1 plants. Moreover, exogenous application of the ethylene precursor 1-aminocyclopropane-1-carboxylate (ACC) induced wild-type levels of resistance in myb72-1, suggesting that MYB72 acts upstream of ethylene in the ISR pathway. Collectively, this study identified the transcriptional regulator MYB72 as a novel ISR signaling component that is required in the roots during early signaling steps of rhizobacteriamediated ISR. www.plant.org on November 2, 2015 -Published by www.plantphysiol.org Downloaded from

Research paper thumbnail of Late heading of perennial ryegrass caused by introducing an Arabidopsis homeobox gene

Plant Breeding, 2004

Perennial ryegrass (Lolium perenne L.) is the most important temperate forage grass species. Unfo... more Perennial ryegrass (Lolium perenne L.) is the most important temperate forage grass species. Unfortunately, the nutritional value of perennial ryegrass declines as maturity progresses, mainly because of a high concentration of poorly digestible compounds in inflorescences. Therefore, the development of forage-type ryegrass varieties with extended vegetative growth is of interest for agriculture. To delay floral transition in perennial ryegrass the Arabidopsis ATH1 gene driven by the maize ubiquitin promoter, the rice actin promoter or the rice OSH1 promoter, respectively was introduced. In ATH1-expressing plants heading was delayed, and in a number of cases the plants never flowered at all. Such non-or late-heading was accompanied by the outgrowth of normally quiescent lateral meristems into extra leaves, resulting in a leafy growth habit. When eventually heading, these plants generally produced a reduced number of inflorescences. These observations suggest that ATH1-mediated delay of heading may be useful to improve fodder quality of perennial ryegrass.