Variation in shoot mortality within crowns of severely defoliated Betula maximowicziana trees in Hokkaido, northern Japan

To clarify mortality patterns of current-year shoots within the crown of Betula maximowicziana Regel after severe insect herbivory in central Hokkaido, northern Japan, we investigated the degree of defoliation, pattern of shoot development, shoot mortality, and leaf tissue-water relations. One hundred current-year long shoots growing in a B. maximowicziana plantation were observed for defoliation and mortality in June 2002. An outbreak of herbivorous insects (Caligula japonica and Lymantria dispar praeterea) occurred in the stand in mid-to-late June, and the monitored shoots were defoliated to various degrees. Within 1 month of defoliation, some of the severely defoliated shoots had produced new leaves on short shoots that had emerged from axillary buds. Stepwise logistic regression revealed that the probability that current-year long shoots would put out axillary short shoots with leaves is closely related to the degree of defoliation. To evaluate the water relations of the leaves, we determined pressure-volume curves for the leaves that survived the herbivorous insect outbreak and the new leaves that emerged after defoliation. The water potential at turgor loss (Ψl,tlp) and the osmotic potential at full turgidity (Ψπ,sat) were higher for the new leaves than for the surviving leaves, indicating a lower ability to maintain leaf cell turgor against leaf dehydration in the new leaves. Of the 100 shoots, 13 died after the emergence of new leaves. Stepwise logistic regression revealed that the probability that the long shoots would die generally increased with the emergence of new leaves, with increasing shoot height. This result suggests that the combined effect of the vulnerability of newly emerged leaves and low water availability, associated with higher shoot positions within the crown, caused shoot mortality. Based on our results, some possible mechanisms for mortality in severely defoliated B. maximowicziana are discussed.