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First published online September 1, 2004
doi: 10.1242/10.1242/dev.01338


Development 131, 4401-4412 (2004)
Published by The Company of Biologists 2004


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Compound leaves: equal to the sum of their parts?

Connie Champagne and Neelima Sinha*

Section of Plant Biology, University of California, 1 Shields Avenue, Davis, CA 95616, USA



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Fig. 1. Compound leaves can be viewed as (A) collections of simple leaves or (B) equivalent to simple leaves. [B redrawn, with permission, from Kaplan (Kaplan, 1975Go); see http://www.schweizerbart.de].

 


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Fig. 2. Comparison of mature leaf form and KNOX1 expression patterns in simple and compound leaves. (A) Amborella trichopoda has simple leaves. (B) KNOX1 proteins accumulate in the shoot apical meristem (SAM) of A. trichopoda, except in the incipient leaf primordium (red asterisk). (C) Oxalis has compound leaves. (D) In Oxalis, KNOX1 proteins accumulate in the SAM and in developing leaves. LP, leaf primordia; P1, primordium 1; M, meristem. Images adapted, with permission, from Bharathan et al. (Bharathan et al., 2002Go).

 


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Fig. 3. The extent of the adaxial domain determines leaflet placement in compound leaves. (A-C) Scanning electron micrographs of vegetative apices. The adaxial domain has been colored pink. (D-F) Mature leaf form. (A) In the developing leaf blade of wild-type tomato, the adaxial domain extends from the base to the tip. (D) The mature tomato leaf has leaflets arranged along the edge of the adaxial domain. (B) The adaxial domain of transgenic antisense PHAN tomato plants is reduced to the tip of the leaf primordium, and these plants often produce cup shaped leaves (E). (C) The adaxial domain of Schefflera actinophylla is restricted to the tip of the developing leaf. (F) Consequently, leaflets are restricted to the tip of the petiole in this plant. m, meristem; P1, P2, P3 and P4, primordia 1, 2, 3 and 4, respectively. Asterisks indicate developing leaflets; red asterisk denotes region forming cup-shaped blade in antisense PHAN tomato plant. Figure adapted, with permission, from Kim et al. (Kim et al., 2003).

 


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Fig. 4. Model of regulatory relationships between KNOX1 genes, leaf genes and hormones in vegetative apices. Genes are shown in red and hormones are shown in blue. Arrows indicate positive regulation and lines with blunt ends indicate negative regulation. Light blue arrows show the path of polar auxin transport. The dotted line designates the incipient leaf primordium. STM, SHOOT MERISTEMLESS; AS1/AS2, ASYMETRIC LEAVES1/2; BP, BREVIPEDICELLUS; KNAT2, KNOTTED-LIKE2 in A. thaliana; LOB, LATERAL ORGAN BOUDARIES; CK, cytokinin; GA, gibberellic acid.

 


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Fig. 5. KNOX1 genes and PHAN genes are expressed in overlapping regions in developing tomato leaves. (A) The KNOX1 gene LeT6 and LePHAN are expressed in the same domain in wild-type tomato leaf primordia. In situ RT-PCR detection of LePHAN expression in a leaf primordium (left) and in situ hybridization with LeT6 in a comparable leaf primordium (right). The small arrows show expression (green fluorescence in left, and purple in right, panel) in leaflet primordia, and the large arrows show expression in the developing vascular trace. (B) Heterozygous Mouse ears (Me/+) mutant tomato has increased levels of LeT6, decreased amounts of LePHAN, and an increased number of leaflets (KNOX1 overexpression phenotype). Homozygous (Me/Me) mutants have even higher levels of LeT6, causing a greater decrease in LePHAN levels. Consequently, the leaves of these plants cannot produce blades (reduced LePHAN expression phenotype). A was adapted, with permission, from and Kim et al. and Janssen et al. (Kim et al., 2003; Janssen et al., 1998Go). (C) The Curl (Cu) phenotype (due to misexpression of LeT6), an antisense LePHAN phenotype (with cup-shaped leaf), and the Cu phenotype in an antisense LePHAN (with cup-shaped leaf) background. The misexpression phenotype of Cu in the antisense LePHAN background is restricted to the region of LePHAN expression.

 





© The Company of Biologists Ltd 2004