Parenchyma fractions drive the storage capacity of nonstructural carbohydrates across a broad range of tree species

Am J Bot. 2022 Apr;109(4):535-549. doi: 10.1002/ajb2.1838. Epub 2022 Apr 8.


Premise: Nonstructural carbohydrates (NSCs) play a key role in tree performance and functioning and are stored in radial and axial parenchyma (RAP) cells. Whether this relationship is altered among species and climates or is linked to functional traits describing xylem structure (wood density) and tree stature is not known.

Methods: In a systematic review, we collated data for NSC content and the proportion of RAP in stems for 68 tree species. To examine the relationships of NSCs and RAP with climatic factors and other functional traits, we also collected climatic data at each tree's location, as well as wood density and maximum height. A phylogenetic tree was constructed to examine the influence of species' evolutionary relationships on the associations among NSCs, RAP, and functional traits.

Results: Across all 68 tree species, NSCs were positively correlated with RAP and mean annual temperature, but relationships were only weakly significant in temperate species and angiosperms. When separating RAP into radial parenchyma (RP) and axial parenchyma (AP), both NSCs and wood density were positively correlated with RP but not with AP. Wood in taller trees was less dense and had lower RAP than in shorter trees, but height was not related to NSCs.

Conclusions: In trees, NSCs are stored mostly in the RP fraction, which has a larger surface area in warmer climates. Additionally, NSCs were only weakly linked to wood density and tree height. Our analysis of evolutionary relationships demonstrated that RAP fractions and NSC content were always closely related across all 68 tree species, suggesting that RAP can act as a reliable proxy for potential NSC storage capacity in tree stems.

Keywords: climates; nonstructural carbohydrates; radial and axial parenchyma; storage; wood density.

Publication types

  • Systematic Review
  • Research Support, Non-U.S. Gov't

MeSH terms

  • Carbohydrates
  • Magnoliopsida*
  • Phylogeny
  • Wood
  • Xylem*


  • Carbohydrates