Genetic and environmental variation in spring and autumn phenology of biomass willows (Salix spp.): effects on shoot growth and nitrogen economy.

Six commercial willow (Salix spp.) varieties were examined to investigate the effects of genotype and environment on spring and autumn phenology and the relationships between phenology, shoot growth and leaf nitrogen (N) retranslocation. The willows were field-grown under different irrigation and fertilization in central Sweden. Two independent data sets of bud-burst, leaf unfolding duration, growth cessation and the timing of leaf abscission were assessed, and the biomass and leaf N data from the end of the first cutting cycle were used. Specific hypotheses were that (1) spring phenology has a greater effect on the shoot biomass production than autumn phenology; (2) later bud-burst is associated with more rapid leaf unfolding; (3) the timing of leaf abscission has a greater effect on the shoot biomass production than height growth cessation; and (4) later leaf fall is associated with poorer leaf N retranslocation. Bud-burst date varied by 19 and 39 days in the 2 years and leaf unfolding duration varied by 13 and 38 days. Growth cessation varied by 2.5 weeks and completion of leaf abscission (> 90% of leaves shed) by more than 3 weeks between the genotypes and treatments. Bud-burst date was inversely correlated with leaf unfolding duration (R(2) = 0.96). Significant effects of the duration of leafy period (bud-burst to leaf abscission) and bud-burst date on shoot growth were found. Delayed growth cessation and leaf abscission were generally associated with a greater biomass production, but especially the relationship between growth cessation and biomass was weak. The results show that the timing of bud-burst and leaf abscission is more important for willow biomass production than growth cessation. Delayed leaf abscission has a negative effect on leaf N retranslocation and increases the N losses. The results have implications for the breeding of perennial energy crops.