The fluid in the coelomata contains coelomocyte cells that defend the animals against parasites and infections. In some species coelomocytes may also contain a respiratory pigment – red hemoglobin in some species, green chlorocruorin in others (dissolved in the plasma) – and provide oxygen transport within their segments. Respiratory pigment is also dissolved in the blood plasma. Species with well-developed septa generally also have blood vessels running all long their bodies above and below the gut, the upper one carrying blood forwards while the lower one carries it backwards. Networks of capillaries in the body wall and around the gut transfer blood between the main blood vessels and to parts of the segment that need oxygen and nutrients. Both of the major vessels, especially the upper one, can pump blood by contracting. In some annelids the forward end of the upper blood vessel is enlarged with muscles to form a heart, while in the forward ends of many earthworms some of the vessels that connect the upper and lower main vessels function as hearts. Species with poorly developed or no septa generally have no blood vessels and rely on the circulation within the coelom for delivering nutrients and oxygen.