Plants load sugars from photosynthesizing leaves into the phloem of exporting veins either ''apoplastically'' (by using H ؉ ͞sucrose symporters) or ''symplastically'' (through plasmodesmata). The ability to regulate photosynthesis in response to the light environment was compared among apoplastic loaders (pea and spinach) and symplastic loaders (pumpkin and Verbascum phoeniceum). Plants were grown under low light (LL) or high light (HL) or transferred from LL to HL. Upon transfer, pea and spinach upregulated photosynthesis to the level found in HL-acclimated plants, whereas up-regulation in pumpkin and V. phoeniceum was limited. The vein density of pea and spinach was the same in HL and LL. Although spinach did not exhibit anatomical or ultrastructural acclimation to the light environment, in pea, wall invaginations in minor vein companion (transfer) cells were more extensive in HL. Furthermore, upon transfer from LL to HL, these invaginations increased in mature pea leaves. Foliar starch levels in mature leaves of plants transferred from LL to HL were not greater than in HL-acclimated leaves of either apoplastically loading species. In the symplastic loaders, plasmodesmatal frequency per loading cell did not vary with treatment, but vein density and thus total plasmodesmatal frequency were higher in HL. Upon transfer of symplastic loaders, however, vein density remained low, and starch levels were higher than in HL; the incomplete acclimation of photosynthesis upon transfer is thus consistent with a carbon export capacity physically limited by an inability to increase vein and plasmodesmatal density in a mature leaf. apoplastic loading ͉ leaf vein density ͉ photosynthesis ͉ symplastic loading ͉ transfer cells P lants adjust their capacity to carry out photosynthesis in response to the demand for photosynthetic products. This balance between source (mature, photosynthesizing leaves) and sinks (growing, metabolizing, and storing tissues) is continuously adjusted in response to environmental and developmental cues (1, 2). When sink activity is high during rapid growth, photosynthesis rates are high, whereas photosynthesis is typically down-regulated when sink activity is lowered [for example, after fruit removal (3) or in response to decreased soil nitrogen availability (4)]. Similarly, photosynthesis is down-regulated in response to external sugar feeding of leaves or plants (5-7), inhibition of sucrose export resulting from overexpression of an apoplastic invertase (8-10), elevated CO 2 levels (11, 12), and inhibition of export by cold-girdling petioles (10, 13). Despite the preponderance of evidence from such manipulative experiments in support of source-sink feedback regulation of photosynthetic capacity, there has been no examination of the potential role of variation in foliar carbon export features that might influence acclimation of photosynthesis to varying environmental conditions.One of the most profound differences among plant species in terms of carbon export from source tissues is the mode of phloem ...