How do fish breathe?

It’s probable that you know that most of the fishes that inhabit in the Earth breathe due to the presence of gills. However, this is not the only respiratory system present in fishes. In this post, we will review different types of breathing in fishes. 

INTRODUCTION

The respiratory system of fishes have to be adapted to two important limitations of underwater life. On the one hand, the amount of dissolved oxygen is smaller in the water than in the air: at 23ºC, air has 210 ml of oxygen per litre of air, while in freshwater is about 6,6 ml/l and in salt water is 5,3 ml/l. On the other hand, water is much more dense and viscous than air. These limitations explain the adaptations in the breathing of this group of animals.

BREATHING WITH GILLS

The oral cavity of teleostei fishes (modern ray-finned fishes) is communicated with the exterior through the mouth and pharyngeal pouches, lateral openings present in the pharynx in which the gills develop. Thanks to the opercle (or gill cover), a hard structure placed in each side of the head, gills are protected.

The structure of the gills is complex. From branchial arches, curved structures that pierce through pharyngeal pouches in each side of the head, two gill filaments grow forming a V. These filaments produce the gill lamellas, folds of the wall’s filaments with a perpendicular disposition. In each side of the filament, we may find between 10 to 40 lamellas per mm. So, it is in these lamellas where the gas exchange happens because they are a very thin wall of tissue and are well supplied with blood.

Structure of the fishes’ gills (Picture: AS Biology Ms Timms).

So, the oxygenated water that passes through the mouth cross the gills and finally abandon the oral cavity through the opercle, while the blood flows in the opposite direction across lamellas to catch the oxygen.

The larva of many fishes have external gills in each side of the head. In the rest of phases, gills become internal. Fishes with a respiration with gills are hagfishes, lampreys, elasmobranchii and bony fishes.

The hagfish is a species with a breathing with gills (Picture: Natureduca).

BREATHING WITH LUNGS

About 400 bony fish species are known to have the ability of breathing from air, most of them living in freshwater ecosystems. Anyway, most of them have both gills and lungs. These species with the two mechanisms usually use the air in certain occasions:

• When the oxygen level in the water goes down.
• When the temperature increases, so the higher the temperature is, the higher are the oxygen necessities.

Lungfishes (Dipnoi) are among the species with the most advanced system. Their lungs have crests and septums similar to those in the lungs of amphibians. The Australian lungfish (Neoceratodus) can breathe with both gills and a lung. African lungfishes (Protopterus) and the South American lungfish (Lepidosiren) breathe with a complex lung and single gills. These fishes need to compulsorily breathe air, as in the contrary they die.

Lungfishes: Australian lungfish (Neoceratodus forsteri), African lungfish (Protepterus annectens), South American lungfish (Lepidosiren paradoxa) and Devonian lungfish (Dipterus) (Picture: Encyclopaedia Britannica).

OTHER BREATHING MECHANISMS IN FISHES

Many fishes have the capacity of breathing through the skin, specially when they are born because they are so small that they do not have specialised organs. As the animal is growing, gills or/and lungs are developing because the diffusion through the skin is not enough. Anyway, skin may be responsible of a 20% or more of gas exchange in some adult individuals. Others can do it through the mouth, the pharynx, the oesophagus, the intestine or the rectum, as is the case of Hoplosternum.

Hoplosternum has the ability of breathing through the digestive tube (Picture: Free Pet Wallpapers).

Some species have developed cavities beyond the gills, the suprabranchial chambers, which can be filled of air. In other, complex organs developed from a very irrigated branchial arch can be formed and act as a lung. This is the case of the catfish and Electrophorus .

Some fishes have the ability to breathe air without a specific adaptation. This is the case of the American eel (Anguilla rostrata), that cover the 60% of the oxygen requests through the skin and the 40% swallowing air from the atmosphere.

REFERENCES

• Notes of the subject Chordates of the Degree in Biology (University of Barcelona).
• Hickman, Roberts, Larson, l’Anson & Eisenhour (2006). Principios integrales de Zoología. Ed. McGraw Hill (13 ed)
• Hill, Wyse & Anderson (2006). Fisiología animal. Ed. Medica Panamericana

The evolution of amphibians: the conquest of the land

Amphibians were the first group of vertebrates to develop limbs and to be able to leave the water to conquer the land. Even if they are seen as simple and primitive animals by most people, amphibians show a wide diversity of survival strategies which have allowed them to occupy most terrestrial and fresh-water habitats. On this entry we’ll explain some of the aspects related to their evolution, explaining how our ancestors managed to get out of the water.

ORIGIN OF THE AMPHIBIANS

Current amphibians, together with reptiles, birds and mammals are found within the superclass Tetrapoda (“four limbs”), the vertebrate group that abandoned the sea to conquer the land. These first tetrapods were amphibians and they evolved around 395 million years ago during the Devonian period from lobe-finned fish named sarcopterygians (class Sarcopterygii, “flesh fins”) within which we find the coelacanth and the current lungfish.

Specimen of coelacanth (Latimeria chalumnae) a sarcopterygian fish, photo by smerikal.

This group of fish is characterized by its fins which, instead of being formed by rays like in most bony fish, they have a bony base that allowed the subsequent evolution of the limbs of the first amphibians. Within the sarcopterygians, the nearest relatives of the tetrapods are the osteolepiformes (order Osteolepiformes) a group of tetrapodomorph fish that got extinct about 299 million years ago.

Restoration of Eusthenopteron, an extinct osteolepiform, by Nobu Tamura.

ADAPTATIONS TO LIVE ON LAND

The conquest of land was not done from one day to the other; it was possible with the combination of multiple adaptations. Some of the most important characteristics that allowed the first amphibians to leave the water were:

• Evolution of lungs, which are homologous to the gas bladder that allows fish to control its buoyancy. Lungs appeared as an additional way to get oxygen from the air. In fact, there is actually a sarcopterygian family the members of which have lungs to get oxygen from the air, for they live in waters poor on oxygen.

• Dissection of Protopterus dolloi a sarcopteryigian fish with lungs.

• Development of the choanaes, or internal nostrils. While fish present a pair of external nostrils at each side of its snout through which water passes on while swimming, the ancestors of the tetrapods only had one external nostril at each side connected to the internal nostrils, the choanae, which communicated with the mouth. This allowed them to get air through their noses using lung ventilation and this way to smell outside of water.
• Apparition of the quiridium-like limb. The quiridium is the tetrapod’s most basic characteristic. This limb is known for having the differentiated parts: the stylopodium (one bone, the humerus or the femur), the zeugopodium (two bones, the radius or tibia and ulna or fibula) and the autopodium (fingers, hands, toes and feet). While the stylopodium and zeugopodium derived from the sarcopterygian’s fins, the autopodium is a newly-evolved structure exclusive from tetrapods.

Simplified drawing of the structure of the quiridium, by Francisco Collantes.

In short, the relatives of the osteolepiformes developed the tetrapod’s typical characteristics before ever leaving water, because they probably lived in brackish, shallow waters, poor in oxygen and that dried out quickly and often.

THE FIRST AMPHIBIANS

Probably the creature known as Tiktaalik is the closest animal to the mid-point between the osteolepiformes and the amphibians. The first recorded amphibians were labyrinthodonts meaning that their teeth had layers of dentin and enamel forming a structure similar to a maze.

Cross-section of a labyrinthodont tooth, form "On the Genesis of Species", by St. George Mivart.

There were four main groups of primitive amphibians, each characterized by: a group that includes the first animals that were able to get out of water, a second group which contains the ancestors of the amniotes (reptiles, birds and mammals) and two more groups, both candidates to be the ancestors of modern amphibians.

Order Ichthyostegalia

Ichthyostegalians were the first tetrapods to be able to leave the water. They appeared at the late Devonian period and they were big animals with large wide heads, short legs and an aquatic or semi aquatic lifestyle (they probably were pretty clumsy on land). They moved around using mainly their muscular tail with rays similar to that of fish.

Fossil and restoration of Tiktaalik. Photo by Linden Tea.

Similarly to current amphibians, they presented a lateral line (sensory organ that allows fish to detect vibrations and movement underwater) and were able to breathe through their skin (they lost the cosmoid scales of their ancestors). Also, the eggs were laid in the water, from which the tadpoles emerged and later on, they suffered a metamorphosis process to become adults just like current amphibians. Subsequently ichthyostegalians gave rise to the rest of amphibian groups.

Skeletons of Ichthyostega and Acanthostega, two typical ichthyostegalians.

Clade Reptiliomorpha

Reptiliomorphs were the ancestors of amniotes and appeared about 340 million years ago. Most of them were usually large and heavy animals, which presented more advanced adaptations to live on land (laterally-placed eyes instead of dorsally-placed ones and a knobby more impervious skin). Even though, reptiliomorphs still laid their eggs in the water and had larval-stages with gills. It wouldn’t be until the late Carboniferous period when the first amniotes (animals that could lay their eggs on dry land) would emancipate completely from water.

Mounted skeleton of Diadectes a large herbivorous reptiliomorph from the American Museum of Natural History, photo by Ghedoghedo.

Order Temnospondyli

This group is one of the possible candidates to being the ancestors of modern amphibians. This is the most diverse group of primitive amphibians and it survived until the early Cretaceous period, about 120 million years ago. The temnospondyls varied greatly in shape, size and lifestyle.

Restoration of Eryops megacephalus a large temnospondylian predator, by Dmitry Bogdanov.

Most of them were meat-eaters, but some were terrestrial predators, some were semi aquatic and some had returned completely to water. Even though, all species had to return to water to breed for the fertilization was external; while the female was laying clutches of eggs in the water, the male released the sperm over them.

Mounted skeleton of Koskinonodon a 3 metres long temnospondyl, from the American Museum of Natural History, photo by Lawrence.

Within the temnospondyls we can find some of the biggest amphibians that ever lived, such as Prionosuchus, with an estimated length of 4,5 meters and about 300 kilograms of weight. Also, even though their skin was not covered with scales, it wasn’t completely smooth like in modern amphibians.

Restoration of Prionosuchus by Dmitry Bogdanov.

It is believed that this group could be the sister-taxon of modern amphibians, even though there’s one last group which could be a candidate to that post.

Order Lepospondyli

Lepospondyls were a small group of primitive animals which appeared at the early Carboniferous and disappeared at the late Permian period. Even though lepospondyls were not as numerous and smaller than the temnospondyls, they presented a wide range of body shapes and adaptations.

Restoration of Diplocaulus magnicornis, of about 1 metre long was the biggest of all lepospondyls, by Nobu Tamura.

The first lepospondyls looked superficially like small lizards, but subsequently lots of groups suffered processes of limb reduction or loss.

Restoration of Pelodosotis, an advanced lepospondyl, by Dmitry Bogdanov.

The relationship of the lepospondyls with the rest of tetrapods isn’t very clear. Different hypothesis go from some authors arguing that they are a group separated from the labyrinthodonts, some thinking that they are the ancestor of current amphibians and reptiles, and some even saying that they are the ancestors of only a portion of modern amphibians.

Restoration of Lysorophus, a Permian lepospondyl, by Smokeybjb.

As we can see, the classification of primitive amphibians can be an extremely complex thing. On this entry I tried to make a summary of the most important groups of ancient amphibians and, on the next one, we’ll center on the evolution of modern amphibians, the so-called “lissamphibians”, and we’ll look in more detail all the controversies surrounding these curious animals.

REFERENCES

The following sources have been consulted in the elaboration of this entry:

• Cover photo: Tyler Keillor and Beth Rooney.
• http://www.ncbi.nlm.nih.gov/pmc/articles/PMC1691537/pdf/14667343.pdf
• http://www.biology.ualberta.ca/courses.hp/biol606/papers/Ahlberg+1998.pdf
• http://usf.usfca.edu/fac_staff/dever/tetrapod_review.pdf
• http://icb.oxfordjournals.org/content/early/2007/08/13/icb.icm055.full.pdf+html
• http://www.usfca.edu/fac_staff/dever/tetropodevol.pdf
• http://icb.oxfordjournals.org.sci-hub.org/content/5/2/287