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In contrast to higher animals,plants maintain the capacity of generating new organs throughout their lifespan.Stem cells in shoot meristem provide the source of all above-ground organs.InArabidopsis,shoot stem cell homeostasis is maintained by ho-meodomain(HD)transcription factors in two main regulatory pathways.The WUSCHEL(WUS)gene is transcribed in the orga-nizing center(OC)below the stem cells.WUS protein moves into the stem cells where it directly activates CLAVATA3(CLV3)expression and thus specifies the stem cellidentity.SHOOTMER-ISTEMLESS(STM)is expressed throughout the shoot meristem and represses stem cell differentiation(Aichinger et al.,2012)(Figure 1A).Whether and how these two pathways interact to regulate shoot stem cell homeostasis were not clear until now.