Biochemistry and genetics of ACC deaminase: A weapon to "stress ethylene" produced in plants

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Abstract

1-aminocyclopropane-1-carboxylate deaminase (ACCD), a pyridoxal phosphate-dependent enzyme, is widespread in diverse bacterial and fungal species. Owing to ACCD activity, certain plant associated bacteria help plant to grow under biotic and abiotic stresses by decreasing the level of "stress ethylene" which is inhibitory to plant growth. ACCD breaks down ACC, an immediate precursor of ethylene, to ammonia and a-ketobutyrate, which can be further metabolized by bacteria for their growth. ACC deaminase is an inducible enzyme whose synthesis is induced in the presence of its substrate ACC. This enzyme encoded by gene AcdS is under tight regulation and regulated differentially under different environmental conditions. Regulatory elements of gene AcdS are comprised of the regulatory gene encoding LRP protein and other regulatory elements which are activated differentially under aerobic and anaerobic conditions. The role of some additional regulatory genes such as AcdB or LysR may also be required for expression of AcdS. Phylogenetic analysis of AcdS has revealed that distribution of this gene among different bacteria might have resulted from vertical gene transfer with occasional horizontal gene transfer (HGT). Application of bacterial AcdS gene has been extended by developing transgenic plants with ACCD gene which showed increased tolerance to biotic and abiotic stresses in plants. Moreover, distribution of ACCD gene or its homolog's in a wide range of species belonging to all three domains indicate an alternative role of ACCD in the physiology of an organism. Therefore, this review is an attempt to explore current knowledge of bacterial ACC deaminase mediated physiological effects in plants, mode of enzyme action, genetics, distribution among different species, ecological role of ACCD and, future research avenues to develop transgenic plants expressing foreign AcdS gene to cope with biotic and abiotic stressors. Systemic identification of regulatory circuits would be highly valuable to express the gene under diverse environmental conditions.

Figures

  • TABLE 1 | PGPR containing ACC deaminase mediated protection in response to various types of stresses.
  • FIGURE 1 | Reaction mechanism catalyzed by microbial ACC deaminase. Route I: direct β-H (Hydrogen) abstraction, Route II: Addition of nucleophiles followed by β-H (Hydrogen) abstraction. Modified figure adapted from the source ref. Hontzeas et al. (2006).
  • TABLE 2 | List of microorganism with ACC deaminase activity.
  • TABLE 3 | Distribution of ACC deaminase in domain Eukarya.
  • FIGURE 2 | The regulatory circuits of AcdS gene expression in Pseudomonas putida UW4 and other related bacteria. AcdR, regulatory gene for ACC deaminase; AcdB: encoding for glycerophosphoryl diester phosphodiesterase; LRP, Leucine responsive protein; FNR: fumarate nitrate reductase protein; CRP, c-AMP receptor protein; AcdS, gene for encoding ACC deaminase.
  • FIGURE 3 | A model for AcdS gene regulation in nitrogen fixing Mesorhizobium sp. Expression of AcdS is positively regulated by NifA2 protein which binds to σ54 and switch on transcription of AcdS gene. NifA1 is also required in regulation of AcdS but its role is not well-understood.
  • FIGURE 4 | The phylogentic tree constructed from AcdS gene sequence of different bacterial strains. The sequence data obtained from IMG database of JGI were used. The evolutionary history was inferred using the Neighbor-Joining method. The bootstrap consensus tree inferred from 500 replicates was taken to represent the evolutionary history of the taxa analyzed. Branches corresponding to partitions reproduced in less than 50% bootstrap replicates are collapsed. The evolutionary distances were computed using the Maximum Composite Likelihood method and were in the units of the number of base substitutions per site. The analysis involved 465 nucleotide sequences. All positions containing gaps and missing data were eliminated. There were a total of 958 positions in the final dataset. Label of bacterial strains/species showing values lower than 1% was hidden. Evolutionary analyses were conducted in MEGA6.0.

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Singh, R. P., Shelke, G. M., Kumar, A., & Jha, P. N. (2015). Biochemistry and genetics of ACC deaminase: A weapon to “stress ethylene” produced in plants. Frontiers in Microbiology. Frontiers Media S.A. https://doi.org/10.3389/fmicb.2015.00937

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