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To increase the efficiency further, the blood flows in the opposite direction to the water flow. This counter-current arrangement enables the almost fully utilized water of low oxygen content to come into close contact with venous blood with a low partial pressure of oxygen. At the same time, carbon dioxide diffuses from the blood into the interlamellar space.

Again, the counter current flows maximize the diffusion gradients, which may be enhanced still further by the presence of the enzyme carbonic anhydrase at the gill surface, which converts some of the gaseous CO 2 to carbonic acid. When a fish is inactive, the respiratory apparatus is more than sufficient to supply the necessary amount of oxygen to the blood.

Under such conditions, only a proportion of the secondary lamellae may be utilized for respiration, and the concentration of red blood cells may be reduced. The latter may assist the heart in that it will reduce the viscosity of the blood being pumped through the gill capillaries before then passing via the arteries to the capillaries of the various tissues.

In response to a high energy demand, or a low concentration of dissolved oxygen in the water, the fish can respond in two ways: the blood flow can be increased by opening up further secondary lamellae to increase the effective respiratory area it may be difficult to increase significantly the blood flow rate through the capillaries themselves , and the concentration of red blood corpuscles can be increased to raise the oxygen carrying capacity of the blood per unit volume.

The latter can be achieved by reducing the blood plasma volume e. At the same time, the ventilation rate is increased to bring more water into contact with the gills within a unit time. These reactions are quite adequate to compensate for the normal fluctuations of energy demands of the fish and of dissolved oxygen concentrations in the water. One of the consequences, however, of an increased ventilation rate is that there will be an increase in the amount of toxic substances in the water reaching the gill surface where they can be absorbed.

Because the osmotic pressure of the body fluids of fish is considerably higher than that of freshwater, there will be a continuous influx of water across the surface epithelium and a corresponding loss of ions into the water. These fluxes occur over the whole body surface, but particularly at the gills which are relatively unprotected in this respect.

Elsewhere, fluxes can be relatively high in the fins, but the remainder of the body is protected by a tough epithelium and usually scales , and a covering of mucus. The influx of water is balanced by a copious discharge of urine, from which as much sodium and chloride as possible has been re-absorbed in the kidneys. These organs do not control the osmotic pressure of the internal fluids; such control is exerted by special cells in the gill epithelium, whereby sodium is taken up in exchange for hydrogen ions, and chloride in exchange for bicarbonate.


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We understand that you need your replacement Scion heater core fast, and take pride in our ability to ship most Web orders the same day. The concentration of oxygen in water is lower than air and it diffuses more slowly. The density of the water prevents the gills from collapsing and lying on top of each other, which is what happens when a fish is taken out of water.

Higher vertebrates do not develop gills, the gill arches form during fetal development , and lay the basis of essential structures such as jaws , the thyroid gland , the larynx , the columella corresponding to the stapes in mammals and in mammals the malleus and incus. In bony fish , the gills lie in a branchial chamber covered by a bony operculum branchia is an Ancient Greek word for gills. The great majority of bony fish species have five pairs of gills, although a few have lost some over the course of evolution. The operculum can be important in adjusting the pressure of water inside of the pharynx to allow proper ventilation of the gills, so that bony fish do not have to rely on ram ventilation and hence near constant motion to breathe.

Valves inside the mouth keep the water from escaping. The gill arches of bony fish typically have no septum , so that the gills alone project from the arch, supported by individual gill rays. Some species retain gill rakers.

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Though all but the most primitive bony fish lack a spiracle, the pseudobranch associated with it often remains, being located at the base of the operculum. This is, however, often greatly reduced, consisting of a small mass of cells without any remaining gill-like structure.

Marine teleosts also use gills to excrete electrolytes. The gills' large surface area tends to create a problem for fish that seek to regulate the osmolarity of their internal fluids. Saltwater is less dilute than these internal fluids, so saltwater fish lose large quantities of water osmotically through their gills. To regain the water, they drink large amounts of seawater and excrete the salt.

How Salmon Adjust from Fresh Water to Salt Water, and Back Again

Freshwater is more dilute than the internal fluids of fish, however, so freshwater fish gain water osmotically through their gills. In some primitive bony fishes and amphibians , the larvae bear external gills, branching off from the gill arches. Some amphibians retain the external larval gills in adulthood, the complex internal gill system as seen in fish apparently being irrevocably lost very early in the evolution of tetrapods.

Sharks and rays typically have five pairs of gill slits that open directly to the outside of the body, though some more primitive sharks have six or seven pairs. Adjacent slits are separated by a cartilaginous gill arch from which projects a long sheet-like septum , partly supported by a further piece of cartilage called the gill ray.

The individual lamellae of the gills lie on either side of the septum. The base of the arch may also support gill rakers , small projecting elements that help to filter food from the water. A smaller opening, the spiracle , lies in the back of the first gill slit. This bears a small pseudobranch that resembles a gill in structure, but only receives blood already oxygenated by the true gills.

Most sharks rely on ram ventilation, forcing water into the mouth and over the gills by rapidly swimming forward.