22 research outputs found
Passive phloem loading and long-distance transport in a synthetic tree-on-a-chip
Vascular plants rely on differences of osmotic pressure to export sugars from
regions of synthesis (mature leaves) to sugar sinks (roots, fruits). In this
process, known as M\"unch pressure flow, the loading of sugars from
photosynthetic cells to the export conduit (the phloem) is crucial, as it sets
the pressure head necessary to power long-distance transport. Whereas most
herbaceous plants use active mechanisms to increase phloem concentration above
that of the photosynthetic cells, in most tree species, for which transport
distances are largest, loading seems to occur via passive symplastic diffusion
from the mesophyll to the phloem. Here, we use a synthetic microfluidic model
of a passive loader to explore the nonlinear dynamics that arise during export
and determine the ability of passive loading to drive long-distance transport.
We first demonstrate that in our device, phloem concentration is set by the
balance between the resistances to diffusive loading from the source and
convective export through the phloem. Convection-limited export corresponds to
classical models of M\"unch transport, where phloem concentration is close to
that of the source; in contrast, diffusion-limited export leads to small phloem
concentrations and weak scaling of flow rates with the hydraulic resistance. We
then show that the effective regime of convection-limited export is predominant
in plants with large transport resistances and low xylem pressures. Moreover,
hydrostatic pressures developed in our synthetic passive loader can reach
botanically relevant values as high as 10 bars. We conclude that passive
loading is sufficient to drive long-distance transport in large plants, and
that trees are well suited to take full advantage of passive phloem loading
strategies
Subcellular concentrations of sugar alcohols and sugars in relation to phloem translocation in Plantago major, Plantago maritima, Prunus persica, and Apium graveolens
Sugar and sugar alcohol concentrations were analyzed in subcellular compartments of mesophyll cells, in the apoplast, and in the phloem sap of leaves of Plantago major (common plantain), Plantago maritima (sea plantain), Prunus persica (peach) and Apium graveolens (celery). In addition to sucrose, common plantain, sea plantain, and peach also translocated substantial amounts of sorbitol, whereas celery translocated mannitol as well. Sucrose was always present in vacuole and cytosol of mesophyll cells, whereas sorbitol and mannitol were found in vacuole, stroma, and cytosol in all cases except for sea plantain. The concentration of sorbitol, mannitol and sucrose in phloem sap was 2- to 40-fold higher than that in the cytosol of mesophyll cells. Apoplastic carbohydrate concentrations in all species tested were in the low millimolar range versus high millimolar concentrations in symplastic compartments. Therefore, the concentration ratios between the apoplast and the phloem were very strong, ranging between 20- to 100-fold for sorbitol and mannitol, and between 200- and 2000-fold for sucrose. The woody species, peach, showed the smallest concentration ratios between the cytosol of mesophyll cells and the phloem as well as between the apoplast and the phloem, suggesting a mixture of apoplastic and symplastic phloem loading, in contrast to the herbal plant species (common plantain, sea plantain, celery) which likely exhibit an active loading mode for sorbitol and mannitol as well as sucrose from the apoplast into the phloem
Symplasmic transport and phloem loading in gymnosperm leaves
Despite more than 130 years of research, phloem loading is far from being understood in gymnosperms. In part this is due to the special architecture of their leaves. They differ from angiosperm leaves among others by having a transfusion tissue between bundle sheath and the axial vascular elements. This article reviews the somewhat inaccessible and/or neglected literature and identifies the key points for pre-phloem transport and loading of photoassimilates. The pre-phloem pathway of assimilates is structurally characterized by a high number of plasmodesmata between all cell types starting in the mesophyll and continuing via bundle sheath, transfusion parenchyma, Strasburger cells up to the sieve elements. Occurrence of median cavities and branching indicates that primary plasmodesmata get secondarily modified and multiplied during expansion growth. Only functional tests can elucidate whether this symplasmic pathway is indeed continuous for assimilates, and if phloem loading in gymnosperms is comparable with the symplasmic loading mode in many angiosperm trees. In contrast to angiosperms, the bundle sheath has properties of an endodermis and is equipped with Casparian strips or other wall modifications that form a domain border for any apoplasmic transport. It constitutes a key point of control for nutrient transport, where the opposing flow of mineral nutrients and photoassimilates has to be accommodated in each single cell, bringing to mind the principle of a revolving door. The review lists a number of experiments needed to elucidate the mode of phloem loading in gymnosperms