The effector SJP3 interferes with pistil development by sustaining SHORT VEGETATIVE PHASE 3 expression in jujube.

Jujube witches' broom (JWB) is a phytoplasma disease that causes severe damage to jujube (Ziziphus jujuba) crops worldwide. Diseased jujube plants show enhanced vegetative growth after floral reversion, including leafy flower structures (phyllody) and the fourth whorl converting into a vegetative shoot. In previous research, secreted JWB protein 3 (SJP3) was identified as an inducer of phyllody. However, the molecular mechanisms of SJP3-mediated pistil reversion remain unknown. Here, the effector SJP3 was found to interact with the MADS-box protein SHORT VEGETATIVE PHASE 3 (ZjSVP3). ZjSVP3 was expressed in young leaves and during the initial flower bud differentiation of healthy jujube-bearing shoots but was constitutively expressed in JWB phytoplasma-infected flowers until the later stage of floral development. The SJP3 effector showed the same expression pattern in the diseased buds and promoted ZjSVP3 accumulation in SJP3 transgenic jujube calli. The N-terminal domains of ZjSVP3 contributed to its escape from protein degradation in the presence of SJP3. Heterologous expression of ZjSVP3 in Nicotiana benthamiana produced typical pistil abnormalities, including trichome-enriched style and stemlike structures within the leaflike ovary, which were consistent with those in the mildly malformed lines overexpressing SJP3. Furthermore, ectopic expression of ZjSVP3 directly bound to the zinc finger protein 8 (ZjZFP8) and MADS-box gene SHATTERPROOF 1 (ZjSHP1) promoters to regulate their expression, resulting in abnormal pistil development. Overall, effector SJP3-mediated derepression of ZjSVP3 sustained its expression to interfere with pistil development, providing insight into the mechanisms of pistil reversion caused by JWB phytoplasma in specific perennial woody plant species.

The 'Candidatus phytoplasma ziziphi' effectors SJP1 and SJP2 destabilise the bifunctional regulator ZjTCP7 to modulate floral transition and shoot branching.

Phytoplasmic SAP11 effectors alter host plant architecture and flowering time. However, the exact mechanisms have yet to be elucidated. Two SAP11-like effectors, SJP1 and SJP2, from 'Candidatus Phytoplasma ziziphi' induce shoot branching proliferation. Here, the transcription factor ZjTCP7 was identified as a central target of these two effectors to regulate floral transition and shoot branching. Ectopic expression of ZjTCP7 resulted in enhanced bolting and earlier flowering than did the control. Interaction and expression assays demonstrated that ZjTCP7 interacted with the ZjFT-ZjFD module, thereby enhancing the ability of these genes to directly bind to the ZjAP1 promoter. The effectors SJP1 and SJP2 unravelled the florigen activation complex by specifically destabilising ZjTCP7 and ZjFD to delay floral initiation. Moreover, the shoot branching of the ZjTCP7-SRDX transgenic Arabidopsis lines were comparable to those of the SJP1/2 lines, suggesting the involvement of ZjTCP7 in the regulation of shoot branching. ZjTCP7 interacted with the branching repressor ZjBRC1 to enhance suppression of the auxin efflux carrier ZjPIN3 expression. ZjTCP7 also directly bound to and upregulated the auxin biosynthesis gene ZjYUCCA2, thereby promoting auxin accumulation. Our findings confirm that ZjTCP7 serves as a bifunctional regulator destabilised by the effectors SJP1 and SJP2 to modulate plant development.