3. Composition of brochosomes
 
Brochosomes are extraordinarily stable structures. They are insoluble in water and organic solvents, cold or boiling. They are resistant to heat. Dry masses of brochosomes are highly hydrophobic. Previous studies have indicated that brochosomes contain both protein and lipid components. This conclusion has been based on positive staining by certain histochemical dyes observed using the light microsope (Smith & Littau 1960; Gouranton & Maillet 1968) and electron microscopic observations of brochosomes digested with various enzymes (Gouranton & Maillet 1968). Preliminary results of our ongoing study indicate that ca. 60% of the brochosome skeletons comprise glycine-rich proteins. Some known glycine-rich structural proteins of insects account for the strength of silk (fibroin), egg cases, egg chorion, and flexible cuticle (resilin).
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4. Production of brochosomes
4.1. Malpighian tubules of leafhoppers
4.2. Secretory cells
4.3. Development of brochosomes
4.4. Changes of secretory programs in ontogenesis
4.5. Secretory products other than brochosomes

Brochosomes are produced in special modified segments of the Malpighian tubules of leafhoppers (Cicadellidae). The Malpighian tubules are the main excretory organs of insects, somewhat analagous to kidneys of vertebrates, their primary functions being excretion and ion exchange. However, in some insects the Malpighian tubules or parts of them are modified to produce various materials for external use (cocoons, covers, "spittle", defense). Production of brochosomes is one of the most remarkable cases of acquiring by these organs a secondary function.


4.1. Malpighian tubules of leafhoppers
Leafhoppers have four Malpighian tubules. Each tubule has an inflated middle segment comprising secretory cells which produce brochosomes. Other tubule parts serve excretory function. They display the same structure as the regular Malpighian tubules found in most insects.
 
The drawing shows a typical alimentary canal and the Malpighian tubules of a leafhopper. Brochosomes produced in secretory segments of the tubules are released through the hindgut (arrows).


4.2. Secretory cells
 
The micrograph below shows a cross-section of a brochosome-secreting Malpighian tubule cell of the 1st instar nymph of Cicadella viridis (L.). The cytoplasm displays numerous secretory vacuoles containing developing or mature brochosomes. Vacuoles with mature products release their content into the tubule lumen. Numerous Golgi complexes, where brochosomes originate, are seen in the basal (outer) parts of the cell. Each secretory cells produces a vast number of brochosomes.

Development of brochosomes
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