Abstract:It is currently widely accepted that, aside from green leaves, other plant organs are able to assimilate carbon via the reductive carboxylic acid cycle. Branches, stems, and even roots often have chloroplast-containing cells. The bark of some trees contains up to 750 mg/m2 of chlorophyll. Photosynthetic activity in trees, bushes, and shrubs has been recorded in the living bark of young twigs, branches and main stems, in addition to the living cells of wood, and sometimes even in the pith. Chlorophyll-containing bark and wood tissue are principally subordinated to non-photosynthetic functions, but typically perform effective internal CO2 recycling using CO2 released from respiration. Chloroplast-containing tissues may re-fix 60%-90% of internal CO2 that has respired from woody tissues or has been transported from xylem sap. Many different terms are used to describe "nonfoliar" CO2 fixation in twigs, branches, and stems; including, bark photosynthesis, corticular photosynthesis, chlorenchymal CO2-reduction, stem-internal CO2-fixation, chlorenchymal CO2-refixation, and stem photosynthesis. It is hypothesized that corticular photosynthesis is driven by stem-internal CO2 derived from mitochondrial respiration and maybe also gaseous xylem efflux. Corticular photosynthesis is an essential physiological process in the trunk that positively contributes to total plant carbon, due to its close relationship with stem respiration and sap flow. First, our review summarized the main physiological and ecological functions of corticular photosynthesis. As corticular photosynthesis works in the same way as leaf photosynthesis, photosynthetic carbon reduction is driven by a combination of effective chloroplast structure, essential enzymatic functions, water, light, and carbon dioxide. We showed that these main factors are present in sufficient quantities within the chlorenchymal bark tissues of trees. Corticular photosynthesis, sap flux velocity, and the CO2 concentration of xylem sapwood all influence stem CO2 efflux. Observations of the relationship between sap flux and CO2 efflux may help explain why CO2 efflux changes with stand age or tree size, in addition to differences between similar trees growing in different environments. Second, we described the principal methods used to measure and calculate corticular photosynthesis. It is now evident that standard measurements of CO2 efflux to the atmosphere, such as a flux chamber covering a segment of tree stem to estimate the rate of woody tissue respiration, do not adequately account for internal fluxes in CO2. The new mass balance approach of measuring corticular photosynthesis may provide a more accurate way of estimating the rate of woody tissue respiration. A more complete assessment of internal CO2 fluxes in stems will improve our understanding about the carbon balance of trees. Third, we discuss the problems and challenges associated with the study of corticular photosynthesis. The unpredictability of stem respiration measurements could be reduced by incorporating corticular photosynthesis measurements into the mass balance correction. We propose that the combined approach of using stable carbon isotope tracing, CO2 and O2 micro-sensors, and sap-flow techniques should be used in future so that the fraction of each source of internal CO2 in the stem-and the respective determinant-may be accurately determined. In addition, we propose that the genomic regulatory mechanisms that influence corticular photosynthesis should be investigated to understand this important process at the gene level. Finally, we suggest that it is important to integrate scaling and model-fitting with eddy covariance and remote sensing techniques to improve estimation accuracy at a regional scale.