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Specification of bundle sheath cell fates during maize leaf development: roles of lineage and positional information evaluated through analysis of the tangled1 mutant

Julia P. Jankovsky^1,, Laurie G. Smith² and Timothy Nelson^1,*

¹ Department of Molecular, Cellular and Developmental Biology, Yale University, PO Box 208104, New Haven CT 06520, USA
² Section of Cell and Developmental Biology, University of California, San Diego, La Jolla, CA 92093, USA
Present address: Tufts School of Veterinary Medicine, North Grafton, MA, USA

View larger version (140K): [in a new window]   Fig. 1. tan1 vascular pattern. Cleared leaf sections of (A) wild-type and (B) tan1, showing the alteration in vascular pattern in the tan1 mutant. Paradermal leaf sections of (C) wild-type and (D) tan1 in Spurr’s resin illustrating the irregular arrangement of BS cell clusters around a minor vein in the tan1 mutant. Arrows point to BS cells in wild-type (C) and ectopic BS cell clusters in the tan1 mutant (D). Bar, 10 µm (C,D).

View larger version (113K): [in a new window]   Fig. 2. tan1 vein phenotype. (A,B) Cross sections through wild-type minor veins and (C-H) tan1 minor veins in Spurr’s resin illustrating the ectopic cell clusters observed around veins in the tan1 mutant. Arrows point to BS cells in wild type (A,B) and ectopic BS cell clusters in the tan1 mutant (C-H). Bar, 10 µm.

View larger version (119K): [in a new window]   Fig. 3. Immunolocalizations of BS and M cell-specific antibodies. BS cell-specific localization of the ME antibody in sections of (A) wild type and (C) tan1, including localization in the ectopic cell clusters. (B) M cell-specific localization of the PEPCase antibody in sections of wild type and (D) tan1. Arrows indicate ME localization in wild-type BS cells (A), tan1 mutant ectopic BS cell clusters (C) and PEPCase localization in wild-type BS cells (B) and tan1 mutant ectopic BS cell clusters (D). All tissue was embedded in Paraplast Plus for this experiment. Bar, 10 µm.

View larger version (78K): [in a new window]   Fig. 4. Examples of sector classes. Class I sectors have lateral boundaries either among M cells (A) or include a complete ring of BS cells (B). Class II sectors have boundaries that fractionate the BS cell ring (C). BS cells in an aberrant cell cluster in tan1 were without exception clonally related to at least one of the normal BS cells surrounding the adjacent vein (D, E). Neither D nor E depicts a section through a transverse vein, as determined by examination of serial sections.

View larger version (79K): [in a new window]   Fig. 5. Cell division patterns in tan1 vein formation. Cell divisions present in the formation of wild-type minor veins (A,B) and tan1 minor veins (C,D), where divisions appear much later than in comparable wild-type veins. Arrows indicate procambial strands in wild-type (A,B) and late divisions within procambial strands in the tan1 mutant (C,D). Bar, 10 µm.