Late summer photosynthesis and storage carbohydrates in walnut (Juglans regia L.): Feed-back and feed-forward effects.

The effect of late summer - autumn limitation of phloem export on growth, photosynthesis and storage carbohydrate accumulation, was evaluated in walnut (Juglans regia L.). This was done by girdling current years shoots, with either all or with only a third of the leaves left in place. Nineteen days after girdling, photosynthesis was greatly reduced and after 46 days, it was about 70% lower in both girdling treatments compared to the control (ungirdled shoots). This reduction is consistent with a feed-back effect of an increased carbohydrate content of the leaves. At the end of the experiment (46 days after girdling), the radial growth of girdled shoots was increased at their base but not at their apical part compared to the control. Girdling increased the accumulation of sucrose in the bark at the base of the shoot and of starch in the bark and in the wood of the shoot apical part. The activity of ADP-glucose pyrophosphorylase in wood increased in the apical part of girdled shoots. The results suggest that a high availability of carbohydrates elicits a feed-forward action on the shoot sink size and activity (radial growth and storage carbohydrate accumulation). Further, for the first time in tree wood we found an increased total activity of AGP induced by an increased assimilate availability. Moreover, the results indicated that, in late summer - autumn, CO2 uptake by leaves of the deciduous tree walnut is strongly dependent on export of photosynthates from the crown. Therefore, carbon uptake in this period depends largely on the availability of effective storage sinks where newly produced assimilates can be accumulated.