1. Introduction
The
Classically,
Plant-microbe interactions often lead to the development of defense mechanisms in plant cells. Since reactive oxygen species (ROS) play a pivotal role in the regulation of plant defense mechanisms, extensive experiments were performed to study the relationship between secondary metabolism (phytoalexin production) and the production of ROS in cells transformed with
2. Agrobacterium rhizogenes
rol genes as activators of secondary metabolism
The interest in
The
3. Agrobacterium and ROS
Reactive oxygen species play an important role during plant-pathogen interactions. Avirulent and virulent pathogens elicit ROS accumulation in plant cells with different dynamics, and elicitors of defense responses, often referred to as microbe-associated molecular patterns (MAMPs), also trigger oxidative bursts (Torres et al., 2006). ROS act as executioners of pathogens and host cells by causing a hypersensitive response; they also act as signaling molecules that activate defense mechanisms. To ensure their own survival, pathogens commonly inactivate ROS produced during plant-pathogen interactions. The plant pathogen
4. ROS and secondary metabolism
In some plant cell cultures, ROS are shown to be sufficient for the induction of plant secondary metabolite accumulation, whereas they are not involved in regulation of secondary metabolism in some other plants (reviewed by Zhao et al., 2005). It has been shown that ROS mediate the elicitor-induced accumulation of isoflavonoids in soybean and alfalfa, indole alkaloids in
The involvement of the oxidative burst generated by NADPH oxidase in the process of phytoalexin stimulation is well known (Guo et al., 1998; Jabs et al., 1997). There are, however, several examples of Ca2+-dependent regulation of defense genes, where the NADPH oxidase pathway is not involved (Romeis et al., 2000; Sasabe et al., 2000). It is clear that several different mechanisms regulate secondary metabolism in plants. Although the details of these mechanisms in different regulatory situations are poorly investigated, the general rule postulates that ROS are important inductors of secondary metabolism.
5. Unexpected complicity of related genes: rolC inhibits ROS production and rolB activates ROS degradation
5.1. ROS levels in transformed cells
Fig. 1 presents results indicating inverse relationship between the production of secondary metabolites (anthraquinones) and ROS levels in callus cultures of
The effect of the
5.2. A model in which rolC inhibits NADPH oxidase via CDPK
It is known that particular calcium-dependent protein kinase (CDPK) isoforms could activate stress-induced NADPH oxidases of plants by phosphorylation. For example, potato StCDPK5 induces the phosphorylation of StRBOHB (
By investigating CDPK genes of
Additional experiments showed that
These results led us to postulate that
In a new research, an Arabidopsis CDPK gene was expressed in
5.3. A model for rolB : NADPH oxidase activation leads to the induction of antioxidant defense system
In contrast to
5.4. How do the rol genes stabilize the biosynthesis of secondary metabolites?
In some cases, the effect of secondary metabolism activation mediated by the
It is evident that the
Investigation of such complex processes is a subject of systems biology. Comprehensive study of the regulatory networks involved in the biosynthesis of secondary metabolites by proteomics methods is an exciting and new field of knowledge (Bulgakov et al., 2011). This methodology will be used to unravel the complex mechanisms of the
6. Combined effect of the rolA , B and C genes
The combined action of the
It is clear that the combined actions of the
Let's consider the situations in which the
In addition, these results reveal an interesting analogy between
7. Conclusion
The combination of defense mechanisms, coupled with the effect of ROS suppression described in this chapter, represents a unique case of plant-microbe interactions.
Acknowledgments
This work was supported by a grant from the Russian Foundation for Basic Research, by the Grant Program "Molecular and Cell Biology" of the Russian Academy of Sciences and by a grant, “Leading Schools of Thought,” from the President of the Russian Federation.
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