תפריט נגישות
ניגודיות עדינהניגודיות גבוהה
מונוכרום
הדגשת קישורים
חסימת אנימציה
פונט קריא
סגור
איפוס הגדרות נגישות
להורדת מודול נגישות חינם תפריט נגישותניגודיות עדינהניגודיות גבוההמונוכרוםהדגשת קישוריםחסימת אנימציהפונט קריאסגוראיפוס הגדרות נגישותלהורדת מודול נגישות חינםThe signal transduction events occurring during shoot gravitropism are mediated through amyloplasts sedimentation, reorientation of actin filaments in the endodermis, and differential changes in level and action of auxin, associated with differential growth leading to shoot curvature. Since increase in reactive oxygen species (ROS) was shown to be associated with growth, we examined the possible use of antioxidants in controlling the gravitropic response, via their interaction with events preceding shoot bending. Reoriented snapdragon (Antirrhinum majus L.) spikes and tomato (Solanum lycopersicum cv. MicroTom) shoots showed a visual upward bending after a lag period of 3 or 5 h, respectively, which was inhibited by the antioxidants N-acetyl-cysteine (NAC) and reduced glutathione (GSH). This suggests the involvement of oxidative reactions in the process. The two antioxidants prevented the sedimentation of amyloplasts to the bottom of the endodermis cells following 0.5-5 h of snapdragon shoot reorientation, suggesting that oxidative reactions are involved already at a very early signal perception stage prior to the visual bending. In addition, a differential distribution in favor of the lower shoot side of various oxidative elements, including H2O2 concentrations and activity of the NADPH-oxidase enzyme, was observed during reorientation of snapdragon spikes. Application of the two antioxidants reduced the levels of these elements and abolished their differential distribution across the shoot. On the other hand, the activity of the antioxidative enzyme, superoxide dismutase (SOD), which was not differentially distributed across the shoot, increased significantly following application of the two antioxidants. The auxin redistribution in reoriented shoots was analyzed using transgenic tomato plants expressing the GUS reporter gene under the Aux/IAA4 promoter (a generous gift of M. Bouzayen, France). GUS response, detected in control shoots 4 h after their reorientation in favor of the lower shoot side was completely eliminated in the NAC-treated shoots, in which the bending response was inhibited. These results suggest that the gravitropic-induced asymmetric auxin redistribution precedes shoot bending, and NAC inhibits this response by abolishing the auxin gradient across the shoot. Our microarray analysis of tomato shoots, using the Affymetrix Tomato GeneChip, revealed differential changes in expression of 266 genes, occurring during the initial 0.5-5 h of shoot reorientation prior to bending. The differential changes in expression of auxin-related genes in favor of the lower shoot side occurred already following 0.5 h of reorientation, while those of cell wall-related genes, associated with shoot bending, occurred only 3 h following shoot reorientation. Among the identified genes, 11 genes were related to auxin, and 36 genes were associated with oxidative processes, demonstrating the involvement of ROS in early events of shoot gravitropism. Taken together, our results suggest that ROS mediate the early gravityinduced amyloplast sedimentation required for signal perception, as well as the lateral auxin movement across the shoot, which is necessary for the auxin asymmetric distribution leading to shoot upward bending.
The signal transduction events occurring during shoot gravitropism are mediated through amyloplasts sedimentation, reorientation of actin filaments in the endodermis, and differential changes in level and action of auxin, associated with differential growth leading to shoot curvature. Since increase in reactive oxygen species (ROS) was shown to be associated with growth, we examined the possible use of antioxidants in controlling the gravitropic response, via their interaction with events preceding shoot bending. Reoriented snapdragon (Antirrhinum majus L.) spikes and tomato (Solanum lycopersicum cv. MicroTom) shoots showed a visual upward bending after a lag period of 3 or 5 h, respectively, which was inhibited by the antioxidants N-acetyl-cysteine (NAC) and reduced glutathione (GSH). This suggests the involvement of oxidative reactions in the process. The two antioxidants prevented the sedimentation of amyloplasts to the bottom of the endodermis cells following 0.5-5 h of snapdragon shoot reorientation, suggesting that oxidative reactions are involved already at a very early signal perception stage prior to the visual bending. In addition, a differential distribution in favor of the lower shoot side of various oxidative elements, including H2O2 concentrations and activity of the NADPH-oxidase enzyme, was observed during reorientation of snapdragon spikes. Application of the two antioxidants reduced the levels of these elements and abolished their differential distribution across the shoot. On the other hand, the activity of the antioxidative enzyme, superoxide dismutase (SOD), which was not differentially distributed across the shoot, increased significantly following application of the two antioxidants. The auxin redistribution in reoriented shoots was analyzed using transgenic tomato plants expressing the GUS reporter gene under the Aux/IAA4 promoter (a generous gift of M. Bouzayen, France). GUS response, detected in control shoots 4 h after their reorientation in favor of the lower shoot side was completely eliminated in the NAC-treated shoots, in which the bending response was inhibited. These results suggest that the gravitropic-induced asymmetric auxin redistribution precedes shoot bending, and NAC inhibits this response by abolishing the auxin gradient across the shoot. Our microarray analysis of tomato shoots, using the Affymetrix Tomato GeneChip, revealed differential changes in expression of 266 genes, occurring during the initial 0.5-5 h of shoot reorientation prior to bending. The differential changes in expression of auxin-related genes in favor of the lower shoot side occurred already following 0.5 h of reorientation, while those of cell wall-related genes, associated with shoot bending, occurred only 3 h following shoot reorientation. Among the identified genes, 11 genes were related to auxin, and 36 genes were associated with oxidative processes, demonstrating the involvement of ROS in early events of shoot gravitropism. Taken together, our results suggest that ROS mediate the early gravityinduced amyloplast sedimentation required for signal perception, as well as the lateral auxin movement across the shoot, which is necessary for the auxin asymmetric distribution leading to shoot upward bending.
