Genome-wide identification of Ziziphus jujuba TCP transcription factors and their expression in response to infection with jujube witches’ broom phytoplasma

Phytoplasmas are plant pathogenic bacteria that inhabit the phloem sieve elements in infected plants and lack a cell wall. Infection of the woody perennial Ziziphus jujuba with phytoplasma causes jujube witches’ broom (JWB), a disease that causes heavy crop losses. The TEOSINTE BRANCHED 1/CYCLOIDEA/PCF (TCP) transcription factors (TFs) are plant-specific regulators of plant morphogenesis, development, and defense after phytoplasma infection. While the TCP gene family has been studied in many plants, there is no report on TCPs in jujube. In this study, 21 ZjTCP TFs were identified and bioinformatically analyzed based on the genome of the Dongzao cultivar. Another 2 ZjTCPs were found in the genome of the Junzao cultivar. Physicochemical properties of the ZjTCP proteins were quite varied, indicating possible versatility of functions. ZjTCP proteins ranged from 172 to 590 amino acids (aa), had isoelectric points (pIs) between 5.53 (ZjTCP1) and 9.81 (ZjTCP11), and molecular weights ranging from 19,279.98 (ZjTCP11) to 61,784.17 kDa (ZjTCP19). Interaction network mapping predicted several hubs, such as ZjTCP6, ZjTCP7, ZjTCP8, ZjTCP15, ZjTCP16, and ZjTCP19, among which ZjTCP6 and ZjTCP16 were predicted to function in plant defense and morphogenesis. Spatiotemporal expression analysis of the ZjTCPs showed that ten of the ZjTCP genes were detected after infection with ‘ Candidatus Phytoplasma ziziphi’ . ZjTCP6, ZjTCP7, ZjTCP16, and ZjTCP19 were up-regulated after phytoplasma infection. ZjTCP16 showed the most significant increase in transcript levels, after the emergence of disease symptoms. ZjTCP12, ZjTCP15, and ZjTCP18 were down-regulated after phytoplasma infection. We concluded that ZjTCP6 and ZjTCP16 were most likely regulatory factors with roles during the plant response to jujube witches’ broom.