The transformation of the ovary into a fruit after successful completion of pollination and ferti... more The transformation of the ovary into a fruit after successful completion of pollination and fertilization has been associated with many changes at transcriptomic level. These changes are part of a dynamic and complex regulatory network that is controlled by phytohormones, with a major role for auxin. One of the auxin-related genes differentially expressed upon fruit set and early fruit development in tomato is Solanum lycopersicum AUXIN RESPONSE FACTOR 9 (SlARF9). Here, the functional analysis of this ARF is described. SlARF9 expression was found to be auxin-responsive and SlARF9 mRNA levels were high in the ovules, placenta, and pericarp of pollinated ovaries, but also in other plant tissues with high cell division activity, such as the axillary meristems and root meristems. Transgenic plants with increased SlARF9 mRNA levels formed fruits that were smaller than wild-type fruits because of reduced cell division activity, whereas transgenic lines in which SlARF9 mRNA levels were red...
Climacteric fruit ripening, as it occurs in many fruit crops, depends on a rapid, autocatalytic i... more Climacteric fruit ripening, as it occurs in many fruit crops, depends on a rapid, autocatalytic increase in ethylene production. This agriculturally important process has been studied extensively, with tomato simultaneously acting both as a model species and target crop for modification. In tomato, the ethylene biosynthetic genes ACC SYNTHASE2 (ACS2) and ACS4 are highly expressed during fruit ripening, with a combined loss of both ACS2 and ACS4 activity preventing generation of the ethylene burst necessary for fruit ripening. However, the individual roles and importance of ACS2 and ACS4 have not been determined. In this study, we examined specifically the role of ACS4 by comparing the phenotype of an acs4 mutant firstly with that of the wild-type, and secondly with two novel ripening-inhibitor (rin) mutants. Ethylene production during ripening was significantly reduced in both acs4-1, and rin lines, with rin genotypes showing the weaker ethylene burst. Also i) the time between anthesis and the start of fruit ripening and ii) the time required to progress through ripening were significantly longer in acs4-1 than in the wild type, but shorter than in the strongest rin mutant. The delay in ripening was reflected in the lower expression of ripening-related transcripts during the mature green and light red ripening stages. Furthermore, expression of ACS2 and ACS4 was strongly dependent on a functional RIN gene, while ACS2 expression was largely independent of ACS4. Altogether, we show that ACS4 is necessary for normal progression of tomato fruit ripening and that mutation of this gene may provide a useful means for altering ripening traits.
Many semi-aquatic plants respond to flooding by elongating the shoot to reach the water surface. ... more Many semi-aquatic plants respond to flooding by elongating the shoot to reach the water surface. This response is initiated by accumulation of ethylene in the plant due to decreased gas-exchange and continued ethylene production during submergence. Ethylene biosynthesis is often limited by the availability of 1-aminocyclopropane-1-carboxylic acid (ACC), the precursor of ethylene, synthesized by ACC synthase. Here, is reported the
Dark-grown Arabidopsis seedlings develop an apical hook by differential elongation and division o... more Dark-grown Arabidopsis seedlings develop an apical hook by differential elongation and division of hypocotyl cells. This allows the curved hypocotyl to gently drag the apex, which is protected by the cotyledons, upwards through the soil. Several plant hormones are known to be involved in hook development, including ethylene, which causes exaggeration of the hook. We show that gibberellins (GAs) are also involved in this process. Inhibition of GA biosynthesis with paclobutrazol (PAC) prevented hook formation in wild-type (WT) seedlings and in constitutive ethylene response (ctr)1-1, a mutant that exhibits a constitutive ethylene response. In addition, a GA-deficient mutant (ga1-3) did not form an apical hook in the presence of the ethylene precursor 1-aminocyclopropane-1-carboxylate (ACC). Analysis of transgenic Arabidopsis seedlings expressing a green fluorescent protein (GFP)-repressor of ga1-3 (RGA) fusion protein suggested that ACC inhibits cell elongation in the apical hook by inhibition of GA signaling. A decreased feedback of GA possibly causes an induction of GA biosynthesis based upon the expression of genes encoding copalyl diphosphate synthase (CPS; GA1) and GA 2-oxidase (AtGA2ox1). Furthermore, expression of GASA1, a GA-response gene, suggests that differential cell elongation in the apical hook might be a result of differential GA-sensitivity.
The Plant Journal (1997) 11(6), 1265-1271 A homolog of the Arabidopsis thaliana ERS gene is activ... more The Plant Journal (1997) 11(6), 1265-1271 A homolog of the Arabidopsis thaliana ERS gene is actively regulated in Rumex palustris upon flooding Wim H. Vriezen1"2, Cynthia PE van Rijn1, Laurentius ACJ Voesenek2'* and Celestina Mariani1 1 Department of ...
Thirteen α-expansin genes were isolated from Rumex palustris, adding to the six already documente... more Thirteen α-expansin genes were isolated from Rumex palustris, adding to the six already documented for this species. Five α-expansin genes were selected for expression studies in various organs/tissues of R. palustris, with a focus on roots exposed to aerated or O2-deficient conditions, using real-time RT-PCR. Several cases of differential expression of α-expansin genes in the various root types of R. palustris were documented, and the identity of the dominant transcript differed between root types (i.e., tap root vs. lateral roots vs. adventitious roots). Several genes were expressed differentially in response to low O2. In situ hybridizations showed expansin mRNA expression in the oldest region of the tap root was localized to cells near the vascular cambium; this being the first report of expansin expression associated with secondary growth in roots. In situ hybridization also showed abundant expression of expansin mRNA in the most apical 1 mm of adventitious roots. Such early expression of expansin mRNA in cells soon after being produced by the root apex presumably enables cell wall loosening in the elongation zone of roots. In addition, expression of some expansin mRNAs increased in ‘mature zones’ of roots; these expansins might be involved in root hair formation or in formation of lateral root primordia. The present findings support the notion that large gene families of α-expansins enable flexibility in expression for various organs and tissues as a normal part of plant development, as well as in response to abiotic stress.
Ethylene and gibberellins (GAs) control similar developmental processes in plants. The role of et... more Ethylene and gibberellins (GAs) control similar developmental processes in plants. The role of ethylene is at least in part to regulate the accumulation of DELLA proteins, key regulators of plant growth, which suppress the GA response. To expand our knowledge of ethylene-GA crosstalk and to reveal how the modulation of the ethylene and GA pathways affects global plant growth, the gibberellin-insensitive (gai), ethylene-overproducing 2-1 (eto2-1) double mutant, which has decreased GA signalling (resulting from gai) and increased ethylene biosynthesis (resulting from eto2-1), was characterized. Both single mutations resulted in reduced elongation growth. The double mutant showed synergistic responses in root and shoot growth, in induction of floral transition, and in inflorescence length, showing that crosstalk between the two pathways occurs in different plant organs throughout development. Furthermore, the altered ethylene-GA interactions affected root-shoot communication, as evidenced by an enhanced shoot:root ratio in the double mutant. When compared with both single mutants and the wild type, double mutants had enhanced content of active GA(4) at both the seedling and the rosette stages, and, unlike the gai mutant, they were sensitive to GA treatment. Finally, it was shown that synergistic responses in the double mutant were not caused by elevated ethylene biosynthesis but that, in the light, enhanced sensitivity to ethylene may, at least in part, be responsible for the observed phenotype.
Plant hormones are considered to be important mediators of the fruit developmental signal after p... more Plant hormones are considered to be important mediators of the fruit developmental signal after pollination. The role of phytohormones in tomato (Solanum lycopersicum) fruit set was investigated here. Transcriptome analysis of ovaries was performed using two complementary approaches: cDNA-amplified fragment length polymorphism (AFLP) and microarray analysis. The gene expression profiles obtained suggest that, in addition to auxin and gibberellin, ethylene and abscisic acid (ABA) are involved in regulating fruit set. Before fruit development, many genes involved in biotic and abiotic responses are active in the ovary. In addition, genes involved in ethylene and ABA biosynthesis were strongly expressed, suggesting relatively high ethylene and ABA concentrations before fruit set. Induction of fruit development, either by pollination or by gibberellin application, attenuated expression of all ethylene and ABA biosynthesis and response genes within 24 h. It is proposed that the function of ABA and ethylene in fruit set might be antagonistic to that of auxin and gibberellin in order to keep the ovary in a temporally protected and dormant state; either to protect the ovary tissue or to prevent fruit development before pollination and fertilization occur.
The transformation of the ovary into a fruit after successful completion of pollination and ferti... more The transformation of the ovary into a fruit after successful completion of pollination and fertilization has been associated with many changes at transcriptomic level. These changes are part of a dynamic and complex regulatory network that is controlled by phytohormones, with a major role for auxin. One of the auxin-related genes differentially expressed upon fruit set and early fruit development in tomato is Solanum lycopersicum AUXIN RESPONSE FACTOR 9 (SlARF9). Here, the functional analysis of this ARF is described. SlARF9 expression was found to be auxin-responsive and SlARF9 mRNA levels were high in the ovules, placenta, and pericarp of pollinated ovaries, but also in other plant tissues with high cell division activity, such as the axillary meristems and root meristems. Transgenic plants with increased SlARF9 mRNA levels formed fruits that were smaller than wild-type fruits because of reduced cell division activity, whereas transgenic lines in which SlARF9 mRNA levels were red...
Climacteric fruit ripening, as it occurs in many fruit crops, depends on a rapid, autocatalytic i... more Climacteric fruit ripening, as it occurs in many fruit crops, depends on a rapid, autocatalytic increase in ethylene production. This agriculturally important process has been studied extensively, with tomato simultaneously acting both as a model species and target crop for modification. In tomato, the ethylene biosynthetic genes ACC SYNTHASE2 (ACS2) and ACS4 are highly expressed during fruit ripening, with a combined loss of both ACS2 and ACS4 activity preventing generation of the ethylene burst necessary for fruit ripening. However, the individual roles and importance of ACS2 and ACS4 have not been determined. In this study, we examined specifically the role of ACS4 by comparing the phenotype of an acs4 mutant firstly with that of the wild-type, and secondly with two novel ripening-inhibitor (rin) mutants. Ethylene production during ripening was significantly reduced in both acs4-1, and rin lines, with rin genotypes showing the weaker ethylene burst. Also i) the time between anthesis and the start of fruit ripening and ii) the time required to progress through ripening were significantly longer in acs4-1 than in the wild type, but shorter than in the strongest rin mutant. The delay in ripening was reflected in the lower expression of ripening-related transcripts during the mature green and light red ripening stages. Furthermore, expression of ACS2 and ACS4 was strongly dependent on a functional RIN gene, while ACS2 expression was largely independent of ACS4. Altogether, we show that ACS4 is necessary for normal progression of tomato fruit ripening and that mutation of this gene may provide a useful means for altering ripening traits.
Many semi-aquatic plants respond to flooding by elongating the shoot to reach the water surface. ... more Many semi-aquatic plants respond to flooding by elongating the shoot to reach the water surface. This response is initiated by accumulation of ethylene in the plant due to decreased gas-exchange and continued ethylene production during submergence. Ethylene biosynthesis is often limited by the availability of 1-aminocyclopropane-1-carboxylic acid (ACC), the precursor of ethylene, synthesized by ACC synthase. Here, is reported the
Dark-grown Arabidopsis seedlings develop an apical hook by differential elongation and division o... more Dark-grown Arabidopsis seedlings develop an apical hook by differential elongation and division of hypocotyl cells. This allows the curved hypocotyl to gently drag the apex, which is protected by the cotyledons, upwards through the soil. Several plant hormones are known to be involved in hook development, including ethylene, which causes exaggeration of the hook. We show that gibberellins (GAs) are also involved in this process. Inhibition of GA biosynthesis with paclobutrazol (PAC) prevented hook formation in wild-type (WT) seedlings and in constitutive ethylene response (ctr)1-1, a mutant that exhibits a constitutive ethylene response. In addition, a GA-deficient mutant (ga1-3) did not form an apical hook in the presence of the ethylene precursor 1-aminocyclopropane-1-carboxylate (ACC). Analysis of transgenic Arabidopsis seedlings expressing a green fluorescent protein (GFP)-repressor of ga1-3 (RGA) fusion protein suggested that ACC inhibits cell elongation in the apical hook by inhibition of GA signaling. A decreased feedback of GA possibly causes an induction of GA biosynthesis based upon the expression of genes encoding copalyl diphosphate synthase (CPS; GA1) and GA 2-oxidase (AtGA2ox1). Furthermore, expression of GASA1, a GA-response gene, suggests that differential cell elongation in the apical hook might be a result of differential GA-sensitivity.
The Plant Journal (1997) 11(6), 1265-1271 A homolog of the Arabidopsis thaliana ERS gene is activ... more The Plant Journal (1997) 11(6), 1265-1271 A homolog of the Arabidopsis thaliana ERS gene is actively regulated in Rumex palustris upon flooding Wim H. Vriezen1"2, Cynthia PE van Rijn1, Laurentius ACJ Voesenek2'* and Celestina Mariani1 1 Department of ...
Thirteen α-expansin genes were isolated from Rumex palustris, adding to the six already documente... more Thirteen α-expansin genes were isolated from Rumex palustris, adding to the six already documented for this species. Five α-expansin genes were selected for expression studies in various organs/tissues of R. palustris, with a focus on roots exposed to aerated or O2-deficient conditions, using real-time RT-PCR. Several cases of differential expression of α-expansin genes in the various root types of R. palustris were documented, and the identity of the dominant transcript differed between root types (i.e., tap root vs. lateral roots vs. adventitious roots). Several genes were expressed differentially in response to low O2. In situ hybridizations showed expansin mRNA expression in the oldest region of the tap root was localized to cells near the vascular cambium; this being the first report of expansin expression associated with secondary growth in roots. In situ hybridization also showed abundant expression of expansin mRNA in the most apical 1 mm of adventitious roots. Such early expression of expansin mRNA in cells soon after being produced by the root apex presumably enables cell wall loosening in the elongation zone of roots. In addition, expression of some expansin mRNAs increased in ‘mature zones’ of roots; these expansins might be involved in root hair formation or in formation of lateral root primordia. The present findings support the notion that large gene families of α-expansins enable flexibility in expression for various organs and tissues as a normal part of plant development, as well as in response to abiotic stress.
Ethylene and gibberellins (GAs) control similar developmental processes in plants. The role of et... more Ethylene and gibberellins (GAs) control similar developmental processes in plants. The role of ethylene is at least in part to regulate the accumulation of DELLA proteins, key regulators of plant growth, which suppress the GA response. To expand our knowledge of ethylene-GA crosstalk and to reveal how the modulation of the ethylene and GA pathways affects global plant growth, the gibberellin-insensitive (gai), ethylene-overproducing 2-1 (eto2-1) double mutant, which has decreased GA signalling (resulting from gai) and increased ethylene biosynthesis (resulting from eto2-1), was characterized. Both single mutations resulted in reduced elongation growth. The double mutant showed synergistic responses in root and shoot growth, in induction of floral transition, and in inflorescence length, showing that crosstalk between the two pathways occurs in different plant organs throughout development. Furthermore, the altered ethylene-GA interactions affected root-shoot communication, as evidenced by an enhanced shoot:root ratio in the double mutant. When compared with both single mutants and the wild type, double mutants had enhanced content of active GA(4) at both the seedling and the rosette stages, and, unlike the gai mutant, they were sensitive to GA treatment. Finally, it was shown that synergistic responses in the double mutant were not caused by elevated ethylene biosynthesis but that, in the light, enhanced sensitivity to ethylene may, at least in part, be responsible for the observed phenotype.
Plant hormones are considered to be important mediators of the fruit developmental signal after p... more Plant hormones are considered to be important mediators of the fruit developmental signal after pollination. The role of phytohormones in tomato (Solanum lycopersicum) fruit set was investigated here. Transcriptome analysis of ovaries was performed using two complementary approaches: cDNA-amplified fragment length polymorphism (AFLP) and microarray analysis. The gene expression profiles obtained suggest that, in addition to auxin and gibberellin, ethylene and abscisic acid (ABA) are involved in regulating fruit set. Before fruit development, many genes involved in biotic and abiotic responses are active in the ovary. In addition, genes involved in ethylene and ABA biosynthesis were strongly expressed, suggesting relatively high ethylene and ABA concentrations before fruit set. Induction of fruit development, either by pollination or by gibberellin application, attenuated expression of all ethylene and ABA biosynthesis and response genes within 24 h. It is proposed that the function of ABA and ethylene in fruit set might be antagonistic to that of auxin and gibberellin in order to keep the ovary in a temporally protected and dormant state; either to protect the ovary tissue or to prevent fruit development before pollination and fertilization occur.
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Papers by Wim Vriezen