Due to its high sensitivity and specificity for tumor detection, positron emission tomography (PET) has become a standard and widely used molecular imaging technique. Given the popularity of PET, both clinically and preclinically, its use has been extended to study plants. However, only a limited number of research groups worldwide report PET-based studies, while we believe that this technique has much more potential and could contribute extensively to plant science. The limited application of PET may be related to the complexity of putting together methodological developments from multiple disciplines, such as radio-pharmacology, physics, mathematics and engineering, which may form an obstacle for some research groups. By means of this manuscript, we want to encourage researchers to study plants using PET. The main goal is to provide a clear description on how to design and execute PET scans, process the resulting data and fully explore its potential by quantification via compartmental modeling. The different steps that need to be taken will be discussed as well as the related challenges. Hereby, the main focus will be on, although not limited to, tracing 11CO2 to study plant carbon dynamics.
Respired CO2 in woody tissues can build up in the xylem and dissolve in the sap solution to be transported through the plant. From the sap, a fraction of the CO2 can either be radially diffuse to the atmosphere or be assimilated in chloroplasts present in woody tissues. These processes occur simultaneously in stems and branches, making it difficult to study their specific dynamics. Therefore, an 11C-enriched aqueous solution was administered to young branches of Populus tremula L., which were subsequently imaged by positron emission tomography (PET). This approach allows in vivo visualization of the internal movement of CO2 inside branches at high spatial and temporal resolution, and enables direct measurement of the transport speed of xylem-transported CO2 (vCO2). Through compartmental modeling of the dynamic data obtained from the PET images, we (i) quantified vCO2 and (ii) proposed a new method to assess the fate of xylem-transported 11CO2 within the branches. It was found that a fraction of 0.49 min−1 of CO2 present in the xylem was transported upwards. A fraction of 0.38 min−1 diffused radially from the sap to the surrounding parenchyma and apoplastic spaces (CO2,PA) to be assimilated by woody tissue photosynthesis. Another 0.12 min−1 of the xylem-transported CO2 diffused to the atmosphere via efflux. The remaining CO2 (i.e., 0.01 min−1) was stored as CO2,PA, representing the build-up within parenchyma and apoplastic spaces to be assimilated or directed to the atmosphere. Here, we demonstrate the outstanding potential of 11CO2-based plant-PET in combination with compartmental modeling to advance our understanding of internal CO2 movement and the respiratory physiology within woody tissues.
Generally, tree species load photoassimilates passively into the phloem, while herbaceous species load actively. These phloem loading strategies have implications for phloem sugar concentration and growth potential. Whereas, in previous research, phloem loading identification was performed with 14 C-autoradiography, we suggest 11 C-autoradiography, because of its compatibility with plant-PET (positron emission tomography) scans. Because 11 C-autoradiography has been hardly used in plant sciences so far, it was tested in contrasting plant species: one temperate tree species, Populus tremula L., three tropical tree species, Erythrophleum suaveolens (Guill. & Perr.) Brenan, E. ivorense A. Chev., and Maesopsis eminii Engl., and two herbaceous crop species Solanum lycopersicum L. and S. tuberosum L. Our results confirmed that P. tremula is a passive loader, and Solanum spp. are active loaders. Erythrophleum spp. and young leaves of M. eminii showed the expected passive loading strategy, but the mature leaves of M. eminii showed an uncommon pattern. Images corrected for leaf tissue thickness supported that mature leaves of M. eminii used active phloem loading, which is linked to continuous investment in growth and new leaves, supporting the lower carbon storage levels often observed in tropical tree species. With this study, we demonstrate that 11 C-autoradiography is a powerful tool to acquire detailed tracer distribution in leaves to typify phloem loading strategies in plant species.
Plant studies using the short-lived isotope 11 C to label photosynthate via atmospheric carbon dioxide (CO 2 ), have greatly advanced our knowledge about the allocation of recent photosynthate from leaves to sinks. However, a second source for photosynthesis is CO 2 in the transpiration stream, coming from respiration in plant tissues. Here, we use in vivo tracing of xylem-transported 11 CO 2 to increase our knowledge on whole plant carbon cycling. We developed a new method for in vivo tracing of xylem-transported CO 2 in excised poplar leaves using 11 C in combination with positron emission tomography (PET) and autoradiography. To show the applicability of both measurement techniques in visualizing and quantifying CO 2 transport dynamics, we administered the tracer via the cut petiole and manipulated the transport by excluding light or preventing transpiration. Irrespective of manipulation, some tracer was found in main and secondary veins, little of it was fixed in minor veins or mesophyll, while most of it diffused out the leaf. Transpiration, phloem loading and CO 2 recycling were identified as mechanisms that could be responsible for the transport of internal CO 2 . Both 11 C-PET and autoradiography can be successfully applied to study xylem-transported CO 2 , toward better understanding of leaf and plant carbon cycling, and its importance in different growing conditions.
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