The exoskeleton is produced and modified by the epidermal cell layer, and each cell in the epidermis must have the necessary information for producing and depositing the right amount of the right cuticular components at the right time; some of them will later have to modify the secreted products to give a mature material. The timing of the various events is often hormonally controlled, but the quantitative information on how much to produce must be inherent in individual epidermal cells.
A new exocuticle is produced at each molt. A thin, lipid-rich epicuticle is initially secreted from the epidermal cells and deposited beneath the old cuticle, followed by secretion of a thicker procuticle, consisting of chitin and proteins. To allow growth, the total surface area of the new cuticle is larger than that of the old one, and expansion and stretching of the new cuticle take place during and after emergence from the old cuticle (exuvium). Some exoskeletal regions, such as the head capsule, mouthparts, and spines, may be sclerotized before ecdysis; this will aid emergence from the old cuticle. These regions cannot be further expanded but will keep their pre-ecdysial size and shape. Other exoskeletal regions are soft and pliant at ecdysis and are sclerotized soon after emergence when cuticular expansion is complete; as soon as the sclerotization process has started, these regions are irreversibly locked in their new shape.
Sclerotization not only makes the exoskeleton harder and stiffer, it also makes the proteins inextractable and more resistant to enzymatic digestion. Before sclerotization, the exoskeletal proteins are bound to each other and to chitin by various noncovalent links, such as electrostatic interactions, hydrogen bonds, and hydrophobic interactions. Such links can be weakened by changes in pH and ionic strength, making the cuticle more pliant, because displacements of the cuticular components will be easier. During the sclerotization process the proteins are linked firmly to each other, polymerized sclerotizing material fills the voids between proteins and chitin molecules, the cuticle is dehydrated, and deformations of the material will be more difficult.
The sclerotization precursors, A-acetyldopamine (NADA) and A-P-alanyldopamine (NBAD), are synthesized from tyrosine in the epidermal cells. The tyrosine molecules are transformed by decarboxylation and hydroxylation to dopamine, which is acylated to NADA and NBAD. These precursors are secreted from the epidermal cells into the cuticular matrix, where they encounter enzymes (phenoloxidases), which oxidize them to the corresponding orthoquinones. Oxidases of different types (tyrosinases, laccases, peroxidases) have been reported and characterized from cuticle. The quinones produced are highly reactive; they will react spontaneously with histidine and lysine residues in the matrix proteins, resulting in cross-links between neighboring proteins, and they will also react with each other, resulting in complex phenolic polymer mixtures. Depending on the precise reaction conditions, the exoskeleton may remain colorless, or a lighter or darker brown coloration may appear during sclerotization.
The water content of the exoskeleton decreases during incorporation of the sclerotizing precursors into the matrix, probably from a decrease in the number of positively charged amino acid residues in the cuticular proteins, which makes the matrix proteins less hydrophilic. Exclusion of water from the intracuticular voids from accumulation of polymerized material also presumably contributes to dehydration of the exoskeletal material. Often only the exocuticular layer of the sclerites is sclerotized, but in some insects the sclerotization process continues for extended periods after ecdysis, resulting in sclerotization of parts of the endocuticle, although to a lesser extent than the exocuticle.
Both the loss in cuticular water content and the formation of cross-links between proteins contribute to a stabilization of the exoskeletal material. The amounts of sclerotizing material incorporated into the various exoskeletal regions varies from less than 1% to more than 10% of cuticular dry weight. These differences are assumed to be responsible for most of the variation in hardness and stiffness of the various exoskeletal regions. Exocuticle tends to be harder and more difficult to deform than endocuticle, presumably because of more extensive sclerotization. The endocuticular layer will tend to be compressed when a piece of exoskeleton is bent, whereas the stiffer exocuticle will be little deformed, although it will be in tension.
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