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Interestingly, a special class of target RNAs, termed target mimics (TMs), can in turn trigger miRNA degradation in plants\u2014a process genetically dependent on the F-box protein HAWAIIAN SKIRT (HWS). However, the pairing rules governing effective TM sites remain unclear. Here, we systematically investigate the pairing architectures that enable plant TM-directed miRNA degradation. Using transient expression in Nicotiana benthamiana leaves and validation in stable Arabidopsis thaliana transgenic lines, we demonstrate that effective TM sites contain a central or near-central unpaired region\u2014classified as insertions (I-type), mismatches (X-type), or deletions (D-type)\u2014flanked by complementary segments. I-type sites tolerate considerable variation in bulge size and position, whereas X- and D-types are more constrained. By incorporating these features into a predictive pipeline, we identified endogenous TMs with alternative pairing architectures, including EARLY NODULIN-LIKE PROTEIN5 (ENODL5), which fine-tunes miR159 levels during Arabidopsis floral development in an HWS-dependent manner. Duplex stability and non-coding context also contribute to TM efficacy. Together, our findings establish a robust empirical framework for understanding, designing, and predicting miRNA TMs in plants.<\/jats:p>","DOI":"10.1093\/plcell\/koag265","type":"journal-article","created":{"date-parts":[[2026,9,4]],"date-time":"2026-09-04T11:31:24Z","timestamp":1788521484000},"source":"Crossref","is-referenced-by-count":0,"title":["Expanding the repertoire of microRNA target mimicry in plants"],"prefix":"10.1093","volume":"38","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-9990-3258","authenticated-orcid":false,"given":"Youhong","family":"Fan","sequence":"first","affiliation":[{"name":"State Key Laboratory of Genetics and Development of Complex Phenotypes, Zhangjiang mRNA Innovation and Translation Center, School of Life Sciences, Fudan University , Shanghai 200438 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1895-7578","authenticated-orcid":false,"given":"Guodong","family":"Ren","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Genetics and Development of Complex Phenotypes, Zhangjiang mRNA Innovation and Translation Center, School of Life Sciences, Fudan University , Shanghai 200438 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2026,9,7]]},"reference":[{"key":"2026100604473529200_koag265-B1","doi-asserted-by":"publisher","first-page":"207","DOI":"10.1016\/j.cell.2005.04.004","article-title":"microRNA-directed phasing during trans-acting siRNA biogenesis in 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