title: How does climate influence xylem morphogenesis over the growing season? Insights from long-term intra-ring anatomy in Picea abies.
abstract: During the growing season, the cambium of conifer trees produces successive rows of xylem cells, the tracheids, that sequentially pass through the phases of enlargement and secondary wall thickening before dying and becoming functional. Climate variability can strongly influence the kinetics of morphogenetic processes, eventually affecting tracheid shape and size. This study investigates xylem anatomical structure in the stem of Picea abies to retrospectively infer how, in the long term, climate affects the processes of cell enlargement and wall thickening.
  
  
  
  Tracheid anatomical traits related to the phases of enlargement (diameter) and wall thickening (wall thickness) were innovatively inspected at the intra-ring level on 87-year-long tree-ring series in Picea abies trees along a 900 m elevation gradient in the Italian Alps. Anatomical traits in ten successive tree-ring sectors were related to daily temperature and precipitation data using running correlations.
  
  
  
  Close to the altitudinal tree limit, low early-summer temperature negatively affected cell enlargement. At lower elevation, water availability in early summer was positively related to cell diameter. The timing of these relationships shifted forward by about 20 (high elevation) to 40 (low elevation) d from the first to the last tracheids in the ring. Cell wall thickening was affected by climate in a different period in the season. In particular, wall thickness of late-formed tracheids was strongly positively related to August-September temperature at high elevation.
  
  
  
  Morphogenesis of tracheids sequentially formed in the growing season is influenced by climate conditions in successive periods. The distinct climate impacts on cell enlargement and wall thickening indicate that different morphogenetic mechanisms are responsible for different tracheid traits. Our approach of long-term and high-resolution analysis of xylem anatomy can support and extend short-term xylogenesis observations, and increase our understanding of climate control of tree growth and functioning under different environmental conditions.
authors.affiliation: Università degli Studi di Padova, Dept. TeSAF, Viale dell'Università 16, 35020 Legnaro (PD), Italy.
authors.forename: Daniele
authors.initials: D
authors.lastname: Castagneri
authors.affiliation: Swiss Federal Institute for Forest, Snow and Landscape Research WSL, Zürcherstrasse 111, 8903 Birmensdorf (ZH), Switzerland.
authors.forename: Patrick
authors.initials: P
authors.lastname: Fonti
authors.affiliation: Swiss Federal Institute for Forest, Snow and Landscape Research WSL, Zürcherstrasse 111, 8903 Birmensdorf (ZH), Switzerland.
authors.forename: Georg
authors.initials: G
authors.lastname: von Arx
authors.affiliation: Università degli Studi di Padova, Dept. TeSAF, Viale dell'Università 16, 35020 Legnaro (PD), Italy.
authors.forename: Marco
authors.initials: M
authors.lastname: Carrer
mesh_terms: D002980:Climate; D007558:Italy; D009024:Morphogenesis; D028222:Picea / Q000033:anatomy & histology* / Q000254:growth & development; D012621:Seasons; D052584:Xylem / Q000033:anatomy & histology* / Q000254:growth & development
keywords: Cell size; Norway spruce; cell-wall thickness; climate change; quantitative wood anatomy; secondary growth; tracheid; tree ring; xylogenesis