Invertase inhibitors in potato: towards a biochemical and molecular understanding of cold-induced sweetening.

Invertase inhibitors classified as cell wall/apoplastic and vacuolar belonging to the pectin methylesterase family, play a major role in cold-induced sweetening (CIS) process of potato tubers. The CIS process is controlled at the post-translational level via an interaction between invertase (cell wall/apoplastic and vacuolar) by their compartment-specific inhibitors (cell wall/apoplastic and vacuolar). Invertase inhibitors have been cloned, sequenced and functionally characterized from potato cultivars differing in their CIS ability. The secondary structure of the invertase inhibitors consisted of seven alpha-helices and four conserved cysteine residues. The well-conserved three amino acids i.e. Pro-Lys-Phe are known to interact with invertase. Location of the genes encoding cell wall/apoplastic and vacuolar invertase inhibitors on potato chromosome number twelve in a tandem orientation without any intervening genes suggest their divergence into the cell wall and vacuole forms following the event of gene duplication. Under cold storage conditions, the vacuolar invertase inhibitor gene showed developmentally regulated alternative splicing and produce hybrid mRNAs which were the result of mRNA splicing of an upstream region of vacuolar invertase inhibitor gene to a downstream region of the apoplastic invertase inhibitor gene. Transgenic potato tubers overexpressing invertase inhibitors resulted in decreased invertase activity, low reducing sugars and improved processing quality making invertase inhibitors highly potential candidate genes for overcoming CIS. Clustered regularly interspaced short palindromic repeats (CRISPR)/CRISPR-associated protein 9 (Cas9) gene-editing technology offers transgene-free breeding for developing CIS resistant potato cultivars. Moreover, the post-transcriptional regulation of invertase inhibitors during cold storage can be warranted. This review summarizes progress and current knowledge on biochemical and molecular approaches used for the understanding of invertase inhibitors with special reference to key findings in potato.