Cytokinin-promoted secondary growth and nutrient storage in the perennial stem zone of Arabis alpina

Plant J. 2021 Mar;105(6):1459-1476. doi: 10.1111/tpj.15123. Epub 2021 Feb 1.


Perennial plants maintain their lifespan through several growth seasons. Arabis alpina serves as a model Brassicaceae species to study perennial traits. Lateral stems of A. alpina have a proximal vegetative zone with a dormant bud zone and a distal senescing seed-producing inflorescence zone. We addressed how this zonation is distinguished at the anatomical level, whether it is related to nutrient storage and which signals affect the zonation. We found that the vegetative zone exhibits secondary growth, which we termed the perennial growth zone (PZ). High-molecular-weight carbon compounds accumulate there in cambium and cambium derivatives. Neither vernalization nor flowering were requirements for secondary growth and the sequestration of storage compounds. The inflorescence zone with only primary growth, termed the annual growth zone (AZ), or roots exhibited different storage characteristics. Following cytokinin application cambium activity was enhanced and secondary phloem parenchyma was formed in the PZ and also in the AZ. In transcriptome analysis, cytokinin-related genes represented enriched gene ontology terms and were expressed at a higher level in the PZ than in the AZ. Thus, A. alpina primarily uses the vegetative PZ for nutrient storage, coupled to cytokinin-promoted secondary growth. This finding lays a foundation for future studies addressing signals for perennial growth.

Keywords: Arabis alpina; cambium; cytokinin; lipid body; perennial; secondary growth; starch.

Publication types

  • Research Support, Non-U.S. Gov't

MeSH terms

  • Arabis / growth & development
  • Arabis / metabolism*
  • Cytokinins / metabolism*
  • Gene Expression Profiling
  • Lipid Metabolism
  • Plant Growth Regulators / metabolism
  • Plant Growth Regulators / physiology
  • Plant Shoots / growth & development
  • Plant Shoots / metabolism
  • Plant Stems / growth & development
  • Plant Stems / metabolism*
  • Starch / metabolism


  • Cytokinins
  • Plant Growth Regulators
  • Starch