Limitations to using phloem sap to assess tree water and nutrient status

Rapid, reliable tools are needed to infer physiological and nutritional health for managing forest systems. Understanding the processes governing tree health is central to the development of these tools. Non-foliar approaches such as the collection of phloem sap reflect processes governing both the use and acquisition of plant water and nutrients at a wide range of temporal (diurnal to seasonal) and spatial (canopy) scales. Despite this, phloem sap is not commonly employed due to an incomplete understanding of transport and post-photosynthetic processes and their effects on chemical concentrations and carbon isotope discrimination. We highlight the need to characterize the influences of storage, remobilization and transport on the concentrations of metabolites to address the time and spatial decoupling of phloem contents to that of environmental stimuli. A conceptual framework is suggested to focus research on key phenomena regarding metabolite transport and highlight significant advantages, misconceptions and limitations to its application.