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doi: 10.1242/10.1242/dev.00466

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Development of the Drosophila mushroom bodies: elaboration, remodeling and spatial organization of dendrites in the calyx

Sijun Zhu1, Ann-Shyn Chiang2 and Tzumin Lee3,*

1 Department of Molecular and Integrative Physiology, University of Illinois, Urbana, IL 61801, USA
2 Department of Life Science, National Tsing Hua University, Hsinchu, 30043, Taiwan
3 Department of Cell and Structural Biology, University of Illinois, Urbana, IL 61801, USA



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  		Fig. 4. Single-cell analysis of dendritic elaboration patterns of three distinct types of MB neurons. Shown are single-cell MARCM clones of {gamma} (A,D), {alpha}'/ß' (B,E) and {alpha}/ß (C,F) neurons. (D-F) Enlarged views of the dendritic areas of the single MB cells shown in (A), (B) and (C), respectively. Arrows indicate the claw-like structures at the ends of most dendritic branches. Genotype: Hs-FLP/+;FRTG13,UAS-mCD8-GFP/FRTG13,tubP-GAL80;GAL4-OK107/+. Scale bars: 20 µm.

 


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  		Fig. 2. Each clonal unit has its characteristic dendritic territory in the adult MB calyx. (A) A diagram of the front view of the adult MB in the right hemisphere. One notional {alpha}/ß neuron is presented. (B) A sagittal view of the MB showing the rough position (a-h) of each confocal image in (C). (C) Serial representative coronal sections of one adult MB calyx in the left hemisphere from posterior to anterior. GFP-positive clones (green) were generated by induction of flip out in newly hatched larvae (NHL). AL, PL, PM and AM represent the anterolateral, posterolateral, posteromedial and anteromedial clonal unit, respectively. Arrows indicate the fourfold structure at the anterodorsal side of the calyx. Genotype: Hs-FLP/+;UAS>rCD2,y+>mCD8-GFP/+;GAL4-OK107/+. Scale bar: 20 µm.

 


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  		Fig. 1. Larval and adult MB calyces show distinct dendritic organization. Shown are representative confocal sections of larval (A-C) and adult (D-F) MB calyces with flip-out Nb clones generated in newly hatched larvae (NHL). Flip-out clones express mCD8-GFP (green), whereas other MB neurons express rCD2 (red). C,F are merged from A and B, and D and E, respectively. ca, calyx; cb, cell bodies; ped, peduncle. Genotype: Hs-FLP/+;UAS>rCD2,y+>mCD8-GFP/+;GAL4-OK107/+. Scale bar: 20 µm.

 


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  		Fig. 3. Various subtypes of MB neurons show different dendritic distribution patterns. (A-F) Mid-coronal confocal sections of adult MB calyces that contain a newly hatched larvae (NHL)-generated, flip-out Nb clone (green). Various subtypes of MB neurons are selectively labeled using GAL4-201Y (A-C) or GAL4-c739 (D-F). Note gaps (arrows in F) present between adjacent GAL4-c739-positive dendritic areas (outlined by broken lines). Mixing of dendrites across clonal boundaries is specifically seen in the upper central subregion (arrowheads in D-F). For a given clone, the primary neurites of {gamma} neurons are localized on the borders (arrows in C) flanking later-derived MB neurites whereas the primary neurites of last-born {alpha}/ß neurons lie in the center (arrowhead in C). (G-I) Dendrites of GAL4-c739-positive {alpha}/ß neurons label the calycal fourfold structure (arrowheads), as revealed on the most anterior coronal confocal section of the adult MB calyx. C,F,I are merged from A and B, D and E, and G and H, respectively. Genotypes: (A-C) Hs-FLP/+;UAS>rCD2, y+>mCD8-GFP/GAL4-201Y; and (D-I) Hs-FLP/+;UAS>rCD2,y+>mCD8-GFP/GAL4-c739. Scale bar: 20 µm.

 


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  		Fig. 7. Schematic models of MB dendritic elaboration/distribution patterns. (A) Frontal view of various representative MB neurons in the anterolateral clone. (B) An oblique section of the MB calyx from posteroventral to anterodorsal surface. The primary neurites of MB neurons are aligned in order on the posterior surface of the calyx such that late-deposited primary neurites are centrally localized within individual clones, as indicated by patterned arrangement of color patches in the curved strip. Broken lines show the junctions between neighboring clonal dendritic fields. The fourfold domain contributed by the dendrites of pioneer {alpha}/ß neurons is outlined by the dash-dotted line. The space between the dotted line and dash-dotted line represents the two- to threefold domain that is exclusively innervated by early-born {alpha}/ß neurons. For simplicity, one subtype of MB dendrites is shown in each clonal unit. Note that late-derived {alpha}/ß dendrites, but not {gamma} dendrites, are well restricted to individual clonal territories. The calyx is normally tilted forward on the medial side. AM, PM, PL and AL represent the anteromedial, posteromedial, posterolateral and anterolateral clonal units, respectively.

 


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  		Fig. 6. Remodeling allows {gamma} neurons to alter dendritic elaboration patterns. (A-C) Single-cell MARCM clones of adult {gamma} neurons homozygous for a babo mutation. The entire calyx (circled by a broken line) was counterstained using mAb nc82 (red) in C. Note that dendrites of mutant {gamma} neurons from different clonal units tangle together at the center of the calyx (arrows). (D-F) Wild-type single-cell MARCM clones of adult {gamma} neurons. (G-I) Wild-type single-cell MARCM clones of larval {gamma} neurons. Note that larval dendrites of various clonal origins elaborate in the same dendritic field (H, I). Genotype: (A-C) Hs-FLP/+;FRTG13,babo[9],UAS-mCD8-GFP,GAL4-201Y/FRTG13,tubP-GAL80; (D-I) Hs-FLP/+;FRTG13,UAS-mCD8-GFP/FRTG13,tubP-GAL80;GAL4-OK107/+. Scale bar: 20 µm.

 


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  		Fig. 5. {alpha}/ß neurons show birth order-dependent dendritic elaboration patterns. Adult single-cell MARCM clones (green) were generated from ~4 hours before white pupa formation to 2 hours after white pupa formation (A,E,I), 2-6 hours after white pupa formation (B,F,J), 6-8 hours after white pupa formation (C,G,K), or later (D,H,L). {gamma} and {alpha}/ß lobes were counterstained using mAb 1D4 (red) in A-D. Note the pioneer ß axonal branch (arrow in A) extends along the dorsal surface of the ß lobe and stops short of the ß lobe end. (E-H) Enlarged views of the dendritic areas of the single-cell clones in A-D. Calycal regions of multiple single-cell/two-cell clones are shown in I-L. The entire MB calyx was counterstained using mAb nc82 (red) in I. Note that pioneer {alpha}/ß neurons project long dendritic processes into the fourfold domain (arrows in E,I), while extending short branches around the calyx base (arrowhead). Genotype: Hs-FLP/+;FRTG13,UAS-mCD8-GFP/FRTG13,tubP-GAL80;GAL4-OK107/+. Scale bars: 20 µm.

 




© The Company of Biologists Ltd 2003