the middle of the visceral mass. The digestive gland, which is equivalent to a vertebrate liver, diverticulates here, as does the pancreas, and both of these empty into the caecum, a pouch-shaped sac where most of the absorption of nutrients takes place. Indigestible food can be passed directly from the stomach to the rectum where it joins the flow from the caecum and is voided through the anus into the mantle cavity. Cephalopods are short-lived, and in mature squid, priority is given to reproduction; the female Onychoteuthis banksii for example, sheds its feeding tentacles on reaching maturity, and becomes flaccid and weak after spawning. Cardiovascular and excretory systems The squid mantle cavity is a seawater-filled sac containing three hearts and other organs supporting circulation, respiration, and excretion. Squid have a main systemic heart that pumps blood around the body as part of the general circulatory system, and two branchial hearts. The systemic heart consists of three chambers, a lower ventricle and two upper atria, all of which can contract to propel the blood. The branchial hearts pump blood specifically to the gills for oxygenation, before returning it to the systemic heart. The blood contains the copper-rich protein hemocyanin, which is used for oxygen transport at low ocean temperatures and low oxygen concentrations, and makes the oxygenated blood a deep, blue color. As systemic blood returns via two vena cavae to the branchial hearts, excretion of urine, carbon dioxide, and waste solutes occurs through outpockets (called nephridial appendages) in the vena cavae walls that enable gas exchange and excretion via the mantle cavity seawater. Buoyancy Unlike nautiloids which have gas-filled chambers inside their shells which provide buoyancy, and octopuses which live near and rest on the seabed and do not require to be buoyant, many squid have a fluid-filled receptacle, equivalent to the swim bladder of a fish, in the coelom or connective tissue. This reservoir acts as a chemical buoyancy chamber, with the heavy metallic cations typical of seawater replaced by low molecular-weight ammonium ions, a product of excretion. The small difference in density provides a small contribution to buoyancy per unit volume, so the mechanism requires a large buoyancy chamber to be effective. Since the chamber is filled with liquid, it has the advantage over a swim bladder of not changing significantly in volume with pressure. Glass squids in the family Cranchiidae for example, have an enormous transparent coelom