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In the distal (red) isthmus mammillae or mammillary knobs are deposited on the surface of the outer membrane in a regular array pattern. The mammillae are proteoglycan-rich and are thought to control calcification. In the shell gland (similar to a mammalian uterus), mineralization starts at the mammillae and around the outermost membrane fibers. The shell gland fluid contains very high levels of calcium and bicarbonate ions. The thick calcified layer of the eggshell

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