Variations in Anatomy, Associations, and Origins of Kranz Tissue

Although the unique tissue required for C4 photosynthesis in nonsucculent plants is often described as being modified leaf parenchyma sheath, which is positioned meaningfully between the mesophyll externally and the vascular tissues internally, the actual range of locations and known associations make that concept untenable. In origin the Kranz tissue develops from procambium as well as ground parenchyma. It is found in stems as well as leaves. In position Kranz tissue can lie in the parenchyma sheath, in the mestome sheath, isolated in the mesophyll, peripherally in some thick leaves, or within the veins. It can be associated with mesophyll only, mesophyll and colorless parenchyma, mesophyll and sclerenchyma, other Kranz tissue and vascular tissues, mesophyll and mestome sheath, mesophyll and phloem, mesophyll and xylem, epidermis, and, finally, mestome sheath and xylem and phloem. The use of the term Kranz is expounded.