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doi: 10.1242/10.1242/dev.00457

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Duplicate FLORICAULA/LEAFY homologs zfl1 and zfl2 control inflorescence architecture and flower patterning in maize

Kirsten Bomblies1, Rong-Lin Wang2, Barbara A. Ambrose3,*, Robert J. Schmidt3, Robert B. Meeley4 and John Doebley1,{dagger}

1 Labortory of Genetics, University of Wisconsin, Madison, WI 53706, USA
2 Syngenta Biotechnology Inc., 3054 Cornwallis Road, Durham, NC 27709, USA
3 Department of Biology and Center for Molecular Genetics, University of California, San Diego, La Jolla, CA 92093, USA
4 Crop Genetics Research, Pioneer-A DuPont Company, 7300 NW 62nd Avenue, Johnston, IA 50131, USA
* Present address: Instituto de Ecologia, Universidad Nacional Autónoma de México (UNAM), AP-Postal 70-275, Coyoacán 04510, México DF, Mexico



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  		Fig. 1. zfl1 and zfl2 gene structures. (A) Alignment of zfl1 and zfl2 proteins deduced from cDNA sequences with RFL (Kyozuka et al., 1998Go), FLO (Coen et al., 1990Go) and LFY (Weigel et al., 1992Go). Repeated leucine residues are indicated by a star. Similar amino acids were identified using BoxShade version 3.1.1. (B) Exon/intron arrangement and sizes (in bp) of zfl1 and zfl2. Five independently derived Mu insertion sites are indicated with triangles and labeled with corresponding mum allele names.

 


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  		Fig. 2. Expression analysis of zfl. (A) 192 bp RT-PCR product and position of the PstI site used to discern zfl1 and zfl2 transcripts. (B) Inverted ethidium bromide-stained gel images of zfl RT-PCR products restricted with PstI, and actin cDNA synthesis and PCR controls. Developmental stages are indicated. Vegetative shoot apical meristem (Veg. SAM) RNA includes the youngest two to three leaf primordia. Higher cycle numbers were used for this tissue because of low actin amplification. Vegetative leaves were collected prior to emergence from the leaf whorl. `Young tassel' RNA was collected at 34 days, just after reproductive transition, while the apex is producing branches and beginning to initiate spikelet pairs. `Older tassel' RNA was collected from inflorescences with differentiated stamens evident in the florets, but prior to tassel emergence. `Young ears' were 3-5 mm long and producing spikelet pairs and spikelets. `Older ears' were 1-1.5 cm long and had differentiated organs visible in their florets. (C-H) zfl expression analysis by mRNA in situ hybridization. (C) Developing ear. (D) Developing tassel. Developing spikelet-pairs (sp) are visible. (E) Male spikelets (s) developing from the spikelet pair meristem (sp). (F) Spikelet meristems (s) and initiating subtending glume primordia. (G) Branching spikelets forming upper (uf) and lower (lf) florets. Arrows indicate glume (gl) and primordia lemma (l). A floral meristem (fm) with stamens and gynoecium apparent is also visible. (H) Later male floret with developing stamen primordia (st), palea (p), lemma, lower florets (lf), lodicules (lo) and glumes (gl).

 


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  		Fig. 3. (A) Autoradiograph of Southern blot with HindIII-digested genomic DNA showing novel bands corresponding to Mu alleles. (B) Autoradiograph of northern blot of total RNA from developing ears probed with a zfl exon 3 PCR product. Signal was detected only in wild type at about 1400 nt (left). The same membrane was stripped and re-probed with maize ubiquitin cDNA (Ub).

 


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  		Fig. 4. Whole plant defects in zfl double mutant plants. (A) Illustration of wild type (left) and double mutant (right). (B) A wild-type tassel. (C) Apex of a zfl double mutant plant showing several `tassel ears'. (D) Diagrammatic illustration of apical region of an individual double mutant plant showing complex axillary structures with husk leaves, multiple ears, and male (yellow) and female (brown) florets often subtended by husk leaves. (E) Wild type (left) and two double mutant (right) ears from sibling plants. Arrowhead indicates a single kernel on a double mutant ear.

 


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  		Fig. 5. Scanning electron microscopy and histology of developing female reproductive organs. (A) Wild-type ear showing developing spikelet pairs. (B) Double mutant ear with normal spikelet-pair initiation. (C) Wild-type spikelet pair showing two upper florets with glume (gl), palea (p), stamen primordia (arrow; st) and a gynoecial ridge (gr) that surrounds the ovule. The lower floret (lf) is visible. (D) Double mutant spikelet pair with upper floret initiating multiple floret meristems subtended by separate paleas (p). (E) Wild-type female florets showing carpels forming silk (si). (F) Double mutant florets generating abnormal organs in aberrant arrangements; ca, a carpel-like organ in the left spikelet. (G) Wild-type floret with a single, fully formed silk (si). (H) Double mutant floret with many silks. (I) Longitudinal section of a wild-type spikelet showing fully formed silk and the carpel surrounding the ovule (ov). (J) Longitudinal section of a double mutant spikelet. Ectopic florets (ef) are visible. Multiple silks arise from multiple carpel layers surrounding an ovule. (K) Double mutant spikelet showing a spiral of organ primordia at the center of one floret (*). (L) A chimeric carpel on a double mutant floret with vegetative outgrowth (v). (M) An early-arising floret replaced by an inflorescence-like structure. (N) A portion of a highly branched early floret of a double mutant plant.

 


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  		Fig. 6. Scanning electron microscopy and histology of developing male reproductive organs. (A) Developing wild-type male spikelet showing the upper (right) and lower floret with stamens (st), a palea (p) and lemma (l). (B) A double mutant male spikelet showing a broken palea (p), abnormal stamens (abst) in the upper floret, extra vegetative organs (*) in both florets, and a small ectopic floret (ef). (C) Longitudinal section of a wild-type male spikelet showing glumes (gl), lemma (l), and two normal stamens (st). (D) A double mutant spikelet showing glume, lemma and an abnormal stamen with a small locule. (E) An indeterminate double mutant spikelet. (F) Whole-mount image of a double mutant floret with two abnormal stamens and overproliferating vegetative organs. (G) Stamens from a double mutant floret showing twisting and vegetative outgrowths (*). (H) A double mutant floret with overproliferating vegetative organs and no stamens.

 


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  		Fig. 7. Quantitative effects associated with the zfl genotype, excluding the double mutant class. Trait values are plotted on the Y axis of each graph, and active (wild-type) zfl copy number is plotted on the X axis. Grey bars represent the range for each genotype. Diamonds in each category are centered on the mean value for the trait within a genotypic class. The width of the diamond is proportional to the number of individuals in each class, and height represents the 95% confidence interval for each class. The left column of graphs shows associations with zfl1, the middle column shows associations with zfl2, and the right column shows effects associated with total active zfl copy number (numbers of active zfl1 and zfl2 in the plant combined). Each graph has the P value for the associated ANOVA indicated in the upper right corner, and a |d/a| ratio in the lower right corner. (A) Flowering time effects were measured by leaf number and days to pollen production. (B) Inflorescence architecture effects, including tassel branch number, and tassel and ear rank.

 

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