Plants have self-protection system in addition to morphological adaptation. Defense response in plants are the result of interaction of many genes in a network. Hormone signaling pathways are targeted by pathogen to disturb and evade plant defense responses. The stress related phytohormone salicylic acid (SA), jasmonic acid (JA) and ethylene (ET) participate in defense responses to mitigate pathogen effect. Plant hormones like abscisic acid (ABA), gibberellic acid (GA), cytokinin (CK), auxins and brassinosteroid (BR) play a critical role in plant-microbe interactions. BR regulates plant growth and defense, there can also be other hormones involved as well such as GA, ET, JA and SA. These different hormone interact to ensure specificity in response.
Brassinosteroids were originally identified as a group of growth-promoting hormones (Grove et al., 1979). Plant interactions with biotrophs, hemibiotrophs and necrotrophic pathogens are modulated by BR. However, BR induce resistance to biotrophs but susceptibility to most hemibiotrophs and necrotrophs (Yu et al., 2018). Plant immune system is able to detect invading microbes and trigger a defensive response against them. At the surface of plant cells, proteins called pattern recognition receptors (PRRs) are able to recognize specific molecules of microbes and pathogens i.e. plant use PRRs to sense conserved microbe- or pathogen-associated molecular patterns (MAMPs/PAMPS). BR signaling pathway has crosstalk with different MAMP/PAMP-triggered immunity (MTI or PTI) pathway at multiple level such as receptor, cytoplasmic or transcriptional levels (Yu et al., 2018). The growth promoting hormones BR have emerged as crucial regulator of the growth-immunity trade-off (Lozano-Duran and Zipfel 2015). BR-activated transcription factor BRASSINAZOLE RESISTANT 1 (BZR1) regulates many defense genes that negatively regulate immune response (Lozano-Duran et al., 2013).
BR signals are perceived by the BR receptor BRASSINOSTEROID INSENSITIVE1 (BRI1) and its coreceptor BRI1-ASSOCIATED KINASE 1 (BAK1) (Yu et al., 2018). BAK1 is a coreceptor for the plant brassinosteroid receptor, the leucine-rich repeat receptor kinase (LRR-RK) BRI1 (Belkhadir et al., 2012). Flagellin, the main protein of the bacterial flagella, elicit defence responses. Flagellin-sensitive2 (FLS2) is involved in flagellin recognition (Gomez-Gomez and Boller 2000). Recognition of flagellin by LRR-RK FLS2 may function in induced immunity by activating defense responses during plant-bacteria interaction.
BAK1 as co-receptor of pattern recognition receptor for both BR signaling and PAMP signaling may modulate the trade-off between plant growth and immunity (Kong et al., 2012). Receptors exist as independent monomer in absence of any ligand. BAK1 promotes BR and flagellin signaling through ligand induced interaction with BRI1 and FLS2. BAK1 is involved in signaling of other PAMPs to induce innate immunity response and also with signaling of unknown endogenous signal that restrict cell death (Gendron and Wang 2007). BAK1 is not only associated with developmental regulation through the plant hormone receptor BRI1 but also has a functional role in PRR-dependent signaling which initiates innate immunity (Chinchilla et al., 2007). BRI1-BAK1and FLS2-BAK1 complexes coexist in cells indicating that BAK1 might mediate the interplay between BR and PAMP signaling (Kong et al., 2012). Furthermore Belkhadir et al. (2012) observed that the synergistic activities of BRs on MAMP responses require BAK1 and that plant steroid homeostasis is critical in establishment of plant immunity. BR signaling modulates plant immunity during plant growth by regulating immune signaling.
In contrast to the view that BR enhance plant innate immunity De Vleesschauwer et al. (2012) reported that Pythium graminicola exploits endogenous BRs as virulence factors and hijacks the host BR machinery to cause disease. P. graminicola coopts the plant BR machinery as a decoy strategy to tap into immune signaling circuitry and interfere with effectual SA- and GA-controlled defences. BR antagonizes SA-mediated root immunity against pathogen P. gaminicola (De Vleesschauwer et al., 2012). It is demonstrated that brassinosteroid-mediated disease resistance (BDR) takes part in plant defense responses independently from SA-mediated defense responses (Nakashita et al., 2003). Albrecht et al. (2012) result, suggested that BR-mediated growth directly antagonizes innate immune signaling. They also observed that activation of BRI1 pathway leads to inhibition of PTI signaling mediated by several PRRs. BR production may contribute to the ability of the pathogen to suppress plant immunity.
BRs and GA undergo crosstalk with plant defense signaling pathways to fine-tune the regulatory role between growth and immunity (De Bruyne et al., 2014; Lozano-Duran and Zipfel 2015). BR is implicated in plant responses to pathogen attack. Exogenously applied BR confers tolerance to plant from a broad range of pathogen infection. BR-induced systemic resistance appears to be independent of the systemic acquired resistance (SAR) activated by necrotizing pathogens or by SAR inducers such as SA. Hence the possible existence of a steroid hormone–mediated disease resistance in BR-treated plant (Yu et al., 2018).
Brassinolide (BL) is a brassinosteroid that regulates plant growth and induce disease resistance in plants. BL has the ability to induce disease resistance in rice plants. In tobacco BL enhances resistance to the viral, bacterial and fungal pathogens (Nakashita et al., 2003). Nakashita et al. (2003) observed BL induces disease resistance in both tobacco and rice, the resistance is systemically induced in tobacco with a mechanism different from known induced resistance. Kemmerling et al. (2007) studied that BAK1 which operates as a coreceptor of BRI1 in brassinolide-dependent plant development, regulates the containment of infection-induced cell death. BAK1-deficient plant develop necrosis upon infection.
Plant receptor kinases may perform dual function for many plant. In tomato, the BRI1 ortholog is required for responses to both BR and peptide hormone systemin which activates the systemic wound responses (Scheer et al., 2003). Such dual function along with several LRR-RLK genes in plant genome (over 230 in Arabidopsis) allows plant to respond to internal and external signals (Gendron and Wang 2007). Programme cell death may act as a barrier to biotrophic pathogens but defense strategies that result into cell death make plant more susceptible to necrotrophic pathogen and therefore must be controlled.
See Part VI for further information
Continue…….
References:
Albrecht, C., Boutrot, F., Segonzac, C., Schwessinger, B., Gimenez-lbanez, S., Chinchilla, D., Rathjen, J. P., de Vries, S. C. and Zipfel, C. 2012 Brassinosteroids Inhibit Pathogen-Associated Molecular Pattern-Triggered Immune Signaling Independent of the Receptor Kinase BAK1. PNAS 109(1): 303 – 308
doi/10.1073/pnas.1109921108
Belkhadir, Y., Jaillais, Y., Epple, P., Balsemao-Pires, E., Dangl, J. L. and Chory, J. 2012 Brassinosteroids Modulate the Efficiency of Plant Immune Responses to Microbe-Associated Molecular Patterns. PNAS USA 109(1): 297 – 302
doi/10.1073/pnas.1112840108
Chinchilla, D., Zipfel, C., Robatzek, S., Kemmerling, B., Nurnberger, T., Jones, J. D., Felix, G. and Boller, T. 2007 A Flagellin-Induced Complex of the Receptor FLS2 and BAK1 Initiates Plant Defence. Nature 448(152): 497 – 500
doi: 10.1038/nature05999
De Bruyne, L., Hofte, M. and De Vleesschauwer, D. 2014 Connecting Growth and Defense: The Emerging Roles of Brassinosteroids and Gibberellins in Plant Innate Immunity. Mol. Plant 7(6): 943 – 959
doi.org/10.1093/mp/ssu050
De Vleesschauwer, D., Buyten, E. V., Satoh, K., Balidion, J., Mauleon, R., Choi, I-R., Vera-Cruz, C., Kikuchi, S. and Hofte, M. 2012 Brassinosteroids Antagonize Gibberellin-and Salicylate-Mediated Root immunity in Rice 1[C][W][OA]. Plant Physiol. 158(4): 1833-1846
doi.org/10.1104/pp.112.193672
Gendron, J. M. and Wang, Z-Y. 2007 Multiple Mechanisms Modulate Brassinosteroid Signaling. Curr. Opin. Plant Biol. 10(5): 436 – 441
doi: 10.1016/j.pbi.2007.08.015
Gomez-Gomez, L. and Boller, T. 2000 FLS2: An LRR Receptor-like Kinase Involved in the Perception of the Bacterial Elicitor Flagellin in Arabidopsis. Mol. Cell 5(6): 1003 – 1011
doi.org/10.1016/S1097-2765(00)80265-8
Grove, M. D., Spencer, G. F., Rohwedder, W. K., Mandava, N., Worley, J. F., Warthen, J. R. D., Steffens, G. L., Flippen-Anderson, J. L. and Cook, J. R. C. 1979 Brassinolide a Plant Growth-Promoting Steroid Isolated from Brassica napus Pollen. Nature 281: 216 – 217
doi:10.1038/281216a0
Kemmerling, B., Schwedt, A., Rodriguez, P., Mazzotta, S., Frank, M., Qamar, S. A., Mengiste, T., Betsuyaku, S., Parker, J. E., Mussig, C., Thomma, Bart P. H. J., Albrecht, C., de Vries, S. C., Hirt, H. and Nurnberger, T. 2007 The BRI1-Associated Kinase 1, BAK1 has a Brassinolide-Independent Role in Plant Cell-Death Control. Curr. Biol. 17(13): 1116-1122
doi.org/10.1016/j.cub.2007.05.046
Kong, X., Pan, J., Cai, G. and Li, D. 2012 Recent Insights into Brassinosteroid Signaling in Plants: Its Dual Control of Plant Immunity and Stomatal Development. Mol. Plant 5(6): 1179 – 1181
doi.org/10.1093/mp/sss097
Lozano-Duran, R., Macho, A. P., Boutrot, F., Segonzac, C., Somssich, I. E. and Zipfel, C. 2013 The Transcriptional Regulator BZR1 Mediates Trade-off between Plant Innate Immunity and Growth. Elife, 2, e00983
doi.org/10.7554/eLife.00983.001
Lozano-Duran, R. and Zipfel, C. 2015 Trade-off between Growth and Immunity: Role of Brassinosteroids. Trends Plant Sci. 20(1): 12- 19
doi: 10.1016/j.tplants.2014.09.003
Nakashita, H., Yasuda, M., Nitta, T., Asami, T., Fujioka, S., Arai, Y., Sekimata, K., Takatsuto, S., Yamaguchi, I. and Yoshida, S. 2003 Brassinosteroid Function in a Broad Range of Disease Resistance in Tobacco and Rice. Plant J. 33(5): 887 – 898
doi: 10.1046/j.1365-313x.2003.01675.x
Scheer, J. M., Pearce, G. and Ryan, C. A. 2003 Generation of Systemin Signaling in Tobacco by Transformation with the Tomato Systemin Receptor Kinase Gene. PNAS USA 100(17): 10114 – 10117
doi: 10.1073/pnas.1432910100
Yu, M-H., Zhao, Z. Z. and He, J-X. 2018 Brassinosteroid Signaling in Plant-Microbe Interactions. Int. J. Mol. Sci. 19(12): 4091
doi.org/10.3390/ijms19124091
BIOLOGICAL PROCESSES- A NATURE'S GIFT 