Issue 92 / March - April 2013
Extraordinary Blood Circulation in Crocodiles
By examining the heart of a crocodile, researchers have discovered how it is that an air-breathing land animal can manage to glide through murky waters for several hours without the need to surface.
There is that one scene in documentaries that we often come across on television: Crocodiles gliding gracefully inside the water, waiting for hours for the perfect time to pounce and snatch their prey from their necks and into the water. But how do these animals manage to stay underwater for almost two hours without surfacing for air even though they, just like human beings and other land animals, live on pulmonary respiration and are in need of the free oxygen in the air?
A member of the reptiles class, crocodiles do not have gill nor can they have skin respiration since their skin is covered with a thick and airtight keratin armor. Just as every living organism are provided with a suitable anatomic and physiological character for their survival, crocodiles are also granted with a system that facilitates their long stay in the water.
Crocodiles are bestowed with a special heart anatomy different to other reptiles like lizards, tortoises, and snakes. The hearts of other reptiles are designed to contain three sections including two atriums and one ventricle. The right atrium, which collects the returned oxygen-deprived (deoxygenated) blood and the left atrium which collects the oxygen-rich (oxygenated) blood retrieved from pulmonary arteries of the lung, transports the blood to one common ventricle. Because there is only one ventricle to receive and combine oxygenated and deoxygenated blood, a mixture of less oxygenated blood is pumped to their body. Depending on outside temperature, the body temperature of a reptile increases or decreases. Their metabolism slows down, almost to a halt, while their body temperature decreases when outside temperature drops near or beyond freezing conditions. Hibernation begins as a result. Frogs and reptiles stop hibernating as soon as their body temperature increases depending on the outside temperature when the weather gets warm. These organisms are called cold blooded animals (with variable body temperatures) because of this feature.
The heart of a crocodile is different to other reptiles in that it has four chambers just like birds and mammals. Blood is sent to the lungs for gas exchange from the right, and from the left ventricle it is pumped to the body. Thus the two types of blood do not mix in the heart. However, what is interesting is that blood is mixed as soon as it leaves the heart via a valve (foramen of panizza) placed in between the right and left aorta.
What could be the purpose of blood, which does not normally mix in the heart, mixing through the medium of a hole? Does this opening in between two aortas indicate a flaw? It is understood after some research that this hole in fact is not a flaw or an anomaly; on the contrary, it is a necessity for a metabolism suited perfectly to the lifestyle of the crocodile.
Warm blooded vertebrates like birds and mammals with a four chamber heart have faster metabolic speeds and higher blood pressures. For these organisms can only supply the energy they consume during their daily activities via such a fast metabolism and a high level of oxygen provided with oxygenated blood.
If the metabolism of a crocodile was fast like mammals all throughout the year, it would have to continuously be nourished and use oxygen. Furthermore, because crocodiles do not have much predators, they could have also lead to the extinctions of some species by overpopulating if they featured a faster metabolism. The low ratio of heart-body mass in crocodiles (0.15%) compared to mammals and birds (0.40%-0.50%) cause the movements of crocodiles to be relatively slower. The Almighty, who creates everything with his wisdom, lowers the blood oxygen ratio and the metabolic speed of crocodiles by creating a valve that combines the two aortas. Thus eliminating the possibility of crocodile overpopulation.
Crocodiles have two aortic arches whereas mammals only have a left, and birds have one right aortic arch. The left aortic arch, despite some contact with the returned blood via foramen of panizza, delivers the oxygenated blood towards intestines, stomach, spleen and the liver after receiving it from the left ventricle of the heart. This is because the digestive system of a crocodile requires oxygen-rich blood. Deoxygenated blood while exiting the right ventricle goes towards the pulmonary arteries of the lung for exchange and mixes with oxygenated blood coming from the right aorta, feeding other organs that are instrumental for its slow metabolism.
Under the water, separated oxygen-rich and oxygen-poor blood mixes when exiting the heart and switches route, thus making oxygenated blood vessels start to carry oxygen poor blood. So what is the reason behind this switch in the direction of the bloodstream under the water? See at this point, the extraordinary features of the crocodile blood circulation system kick in. The two anatomical features belonging only to only crocodile hearts is what enables them to stay under water without breathing. Because of little or no lung use under the water, a big portion of the blood stream is diverted away from lungs; therefore oxygen poor blood is pumped back to the body. As one feature of the two, foramen of panizza restricts (does not close) upon signals coming from nostril sensors under the water but expands and remains open on land. The two aortic arches connects with each other via foramen of panizza as soon as they leave the heart but merge completely in the lower parts of the body away from the heart (anastomosis).
The second feature stems from a serrated valve. Refilling of pumped blood is stopped via a passive, thin leaf-shaped valve which is located at the tip of the pulmonary artery exiting the right ventricle. Thus, one-way direction of blood flow in the heart is maintained. These valves, which carry nodules made of connective tissue, constrict during the dive and blood flow to the lungs is reduced greatly. Therefore blood rejoins the systemic circulation from the right aortic arch.
The blood circulation of crocodiles is similar to birds, mammals, and humans while they are active on land. Oxygen-deprived blood is sent to lungs for gas exchange. The only difference is the turning of the right aorta to the left and the left aorta to the right. Oxygen rich blood not only flows through the left aorta but also through the right aorta via foramen of the panizza as well causing distribution via two channels into the body. However, the foramen of the panizza being open is not sufficient for these two channels to be used. At the same time, the pressure of the blood within the left ventricle needs to be higher as well. This way, high pressure oxygenated blood flows into the right aorta through the opening of the panizza, applying pressure to the valve at the tip of the right aorta to close it in order to prevent the mixing of the oxygen-poor blood into this route. As a result, oxygenated blood gets distributed quickly by each aortic arch without mixing with the used blood. Thus, oxygen-poor and oxygen-rich blood follows the following route on land
* Deoxygenated blood: Body - superior and inferior pulmonary veins - right atrium - right ventricle - lung pulmonary artery - lungs.
* Oxygenated blood: Lung pulmonary vein - left atrium - left ventricle - right aorta and left aorta via panizza valve (both aortas are active) and body.
The opening of the panizza narrows with the help of signals coming from the nostrils when crocodiles submerge. At the same time serrated valves at the tip of pulmonary artery that transports the blood to the lungs also constrict. While this serrated valve is at work, a majority of the blood returning from the body is not sent to the lungs because they are not functioning at the time. This serrated valve also increases the pressure of the right ventricle. This pressure, along with elevated resistance in pulmonary circulation and lowered pressure of systemic circulation, leads to the opening of normal valves at the tip of the left aorta. In the end, the left aorta which normally carries oxygenated blood on land starts carrying oxygen deprived blood, and there is a route switch.
The most beneficial part of this switch is to re-route the deprived blood back to the body via a different route, that is, the left aorta. This by-passes the lungs and prevents time loss. Despite the fact that blood of the left aorta mixes with the oxygenated blood of the right aorta to some degree via the panizza valve, the main function of this opening while submerged is to supply blood flow to the arteries feeding the heart and brain through the transfer of some poor blood from the left aorta into the right aorta; this way vital organs are not left without blood.
Blood returning from the body is not sent to the lungs for gas exchange when crocodiles are under the water. However, existing oxygen in the blood can be delivered to the tissues quickly by a route switch. The amount of bicarbonate ions that is important in the transport of CO2 in the blood increases when oxygen pressure in the tissues drop. Anaerobic respiration of tissues increases. This leads to an increase in lactic acid levels and reduces pH. Eventually, it facilitates the release of oxygen carried by hemoglobin. In the end, oxygen that is bonded with hemoglobin is used more efficiently. In the meantime, the body temperature of a crocodile submerged under water decreases and slows down its metabolism, reducing the need for oxygen. Oxygen stored in the blood can be sufficient up to two hours under the water. However when these reserves are consumed, crocodiles have to resurface to breathe even though it may cost a prey to escape.
Crocodiles can live on land and in the water and adapt to their environments with ease and efficiency thanks to the ponderous working of their mechanisms under water, the change in blood circulation, slow blood flow, reduced body temperature and metabolic speed bestowed upon them. Just like humans in sleep, crocodiles can remain submerged for long periods (4-6 minutes in usual dives; up to 2 hours when pressed) with this perfect system granted to them. Crocodiles use these mechanisms not only when under water, but also while resting or for periods after heavy feeding.