The Role of Hydrogen Peroxide in Controlling Plant Cell-Signaling and Gene-Expression Patterns Related to Stress and Defense Responses

FUTURE WORK

Though many of the signaling pathways for hydrogen peroxide-mediated stress and defense responses have been studied extensively, there exist multiple avenues of additional research that can further clarify the mechanism and modulating factors for these pathways. As one example, future research should attempt to delineate the impact of more endogenous compounds that can strengthen or weaken the wound-response and abiotic stress-response. Nitric oxide has been shown to negatively modulate wound signaling in the tomato plant Lycopersicon esculentum, blocking hydrogen peroxide production and proteinase inhibitor synthesis stimulated by systemin and jasmonic acid (Orozco-Cardenas and Ryan 2001). However, a seemingly opposite trend is present when considering responses to abiotic stressors such as light and drought conditions. In these cases, nitric oxide has been shown to work hand-in-hand with hydrogen peroxide to induce myo-inositol phosphate synthase that confers multiple resistances to abiotic stresses, to reduce stress from drought in marigold plants, and to alter expression of genes in citrus plants to influence acclimation to salinity (Tan et al. 2013; Liao et al. 2012; Tanou et al. 2009). Therefore, it is important to study whether the role of nitric oxide, and other endogenous compounds, in defense responses depends on factors such as specific type of wounding event or stressor by conducting additional research that varies such conditions. As a related suggestion, additional research should focus on studying the well-documented effects of compounds such as glutathione reductase and micronutrients in taxa that have not been studied previously. Learning about variations in mechanism and effect will be crucial for understanding how certain plants can survive in differential environments, and might suggest baseline genetic differences between different species in H₂O₂–mediated signaling pathways.

Another important aspect of future research concerning hydrogen peroxide and its role in plant defense systems involves tracking the molecular basis for how some insects can bypass H₂O₂–based wound responses in plants. Recent studies have suggested that the oxidative burst that occurs after the initial wounding event may actually provide insects with a way to sidestep subsequent plant responses because it allows a timeframe in which the insect can focus on silencing plant defenses while the plant focuses on reducing hydrogen peroxide levels (Kim et al. 2012). Further investigation into the relative timing of the oxidative burst and H₂O₂- scavenging activities will allow researchers to determine the ideal time for insect attempts to bypass natural plant defenses. Such information would allow for the development of protocols involving the use of various exogenous products as added protection for plants. As an example, several micronutrients such as riboflavin and cadmium have been found to boost the strength of H₂O₂–mediated stress and defense responses (Azami-Sardooei et al. 2010; Tamas et al. 2009). A major question is whether such increases in overall magnitude of response can also effectively block insect attempts to silence plant defenses during the scavenging period following oxidative burst.

One last direction for future research is to work towards creating a mathematical model for the level of hydrogen peroxide throughout entire defense or wound-event pathways. Creation of such a model would depend on implementation of multiple trials with different plants and different stressors. Being able to quantitatively assess variations in H₂O₂ from oxidative burst to subsequent signaling pathways would form the basis for more easily discriminating between defense mechanisms. Furthermore, such a real-time model would facilitate wild-type/mutant comparative studies examining the effects of knockout of one or more genes relevant to hydrogen peroxide signaling pathways.

SUMMARY CONCLUSIONS

• Hydrogen peroxide, H₂O_2, is an endogenous molecule in plants that is part of a group of cellular components referred to as reactive oxygen species (ROS) (Orozco-Cardenas, Narvaez-Vasquez and Ryan 2001).
• H₂O₂ activates MAPK cascades and mediates UV-B-induced gene expression, as indicated by down-regulation of UV-induced gene PDF1 in Arabidopsis plants exposed to antioxidants and UV-B light (Lebrun-Garcia et al. 1998; AH Mackerness et al. 1999).
• In wound pathways, H₂O₂ serves as a local signal for hypersensitive cell death and cell wall stiffening, stimulates defense-genes in adjacent cells, and is linked to up-regulation of phenylpropanoid genes expressing phenolic defense compounds (Orozco-Cardenas, Narvaez-Vasquez and Ryan 2001; Lawton et al. 1983).
• The polyamine oxidase gene (NbPAO) is essential for H₂O₂ generation and subsequent hypersensitive response, and may be part of a larger Rboh gene network that includes other respiratory burst oxidase homologs (Yoda et al. 2009; Drerup et al. 2013).
• Many exogenous and endogenous compounds have been implicated in H₂O₂- mediated stress and defense pathways, including several enzyme and micronutrients. Future research should report additional influential structures and clarify known pathways.

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