Wednesday, September 7, 2016

Do You Drink Like A Fish?

Biology concepts – fish osmoregulation, shark osmoregulation, semelparity, iteroparity

The irony of fish drinking is not lost on this café in
the Hotel Portofino at Universal Orlando. What I
really like is the eye patch.
You’d think that fish would never be thirsty; if he needs a drink, he just opens his mouth. But some fish don’t drink a drop! Wouldn’t that be similar to some birds never breathing? Ridiculous.

Fish are good examples of the problems of maintaining proper water and salt concentrations. Some fish live in freshwater, and some in saltwater. These are opposite sides of the same coin when dealing with osmoregulation.

Freshwater fish live in a hypotonic (low salt) environment. The flesh of the fish contains more salt than does the water. Diffusion and osmosis work to equalize salt concentrations in different compartments. Therefore, water will move from the lake or river into the fish’s tissues in order to balance the salt concentrations by osmosis. Salt will not move out of the tissues, since there are molecular mechanisms that work to keep the inside.

Like the kangaroo rat, freshwater fish don’t drink. They do take in water when they eat and move water across their gills, but they don’t take in water just for the water. Even without drinking specifically, freshwater fish take in way more water than they need. Anywhere freshwater contacts a fish cell, water will move inward; this includes the gills, the mouth and gut, and the skin.

In a situation like this, kidney-mediated concentration of urine would be counterproductive; why retain water when water is exactly what you have too much of? Therefore, freshwater fish excrete large amounts of urine. Their kidneys have large glomeruli, which move lots of water into the collecting tubules for excretion.

Saltwater and freshwater fish have different ways of
dealing with salt and water loss and conservation.
Freshwater fish must conserve salt, while saltwater
fish must conserve water. The kidneys play a role,
but so do the chloride cells in the gills.
But if the freshwater fish aren’t drinking, how do they get their salt, which is present in low concentrations in the water? You’d think they would have to be drinking all the time just to collect enough salt.  To get around this, they conserve the salt they ingest through the food they eat. They also take in salts through their gill chloride cells, actively pumping sodium and chloride out of the freshwater and into cells that have a lot of mitochondria (to provide energy to pump the salts). The relatively short collecting tubules of the freshwater fish kidney allow for reuptake of a lot of salt, while excluding almost all the water.

Marine (saltwater) fish have the opposite problem. Their tissues are of much lower salt than they surrounding hypertonic ocean, so osmosis wants to dry them out, sending water out of their bodies. The amount of available drinking water is extremely low - can you imagine dying of dehydration while surrounded by water. Just ask anyone who has survived a shipwreck and prolonged float in the ocean; drinking seawater can be lethal.

However, marine fish must drink all the time in order to keep enough water in their body. Retaining water would be an essential function of marine fish kidneys. They are all fish, but their kidneys work in exactly opposite ways.  Marine kidneys have small or absent glomeruli, so little water is taken out of the blood, but long collecting tubules in order to excrete as much salt as possible.

Drinking a lot of saltwater leaves marine fish with way too much salt; more than their kidneys can get rid of. To aid in salt excretion, they also have chloride cells in their gills. In the opposite fashion of the specialized gill cells of freshwater fish, the chloride cells of saltwater fish actively sequester salts from the blood, and then pump the sodium and chloride out into the seawater.

Sharks have unique ways of maintaining
salt and water. I have no idea of their
mechanisms for pepper regulation.
But sharks are an exception among marine fish. They have a different way to combat high salt concentrations. Remember that osmosis means that water moves from areas of low solute (high water concentration) to areas of high solute (lower water concentration). For many marine fish, this would mean a constant loss of body water to the ocean and quick death by dehydration; much like pouring salt on a slug.

To overcome this movement, sharks produce and retain a huge amount of a chemical called urea; it is one of the soluble wastes that animals normally get rid of. This molecule doesn’t affect the electrical potential that salts create, but increases the solute concentration in the shark’s tissues at levels higher than in the seawater, so water (without the salt) will diffuse into the shark’s body. This is its source of fresh water.

Therefore, sharks are osmoconformers; they maintain an osmotic balance with their environment. If the shark becomes too salty and salt needs to be excreted, it has a salt gland, much like that of birds and reptiles, but the shark’s gland is located in it anus, not near its eyes or nose – that’s a big difference! Taken together, there is no force for movement of water in or out of the shark’s tissues, and the shark remains shark-shaped instead of shriveling or swelling up.

Here is a bullshark caught in the Potomac River.
And you thought that sharks in Washington D.C.
were just in the federal buildings.
An exception to this rule for sharks is the bull shark; it can live in both saltwater and freshwater. Most sharks put into in freshwater would absorb too much water and die of water toxicity. However, the bull shark’s kidneys can adjust to the salinity of the water within a short period of time. Their kidneys will remove less salt and more urea from their blood and tissues and into their urine. They move from being osmoconformers to osmoregulators.

A shark that can live in freshwater; this can present a real problem. There have been many bull shark attacks in rivers and estuaries (video), where people don’t expect to encounter sharks. It is suggested that this behavior and physiology is an adaptation that gives the bull shark a protected nursery for their young, away from predators.

Most fish are stenohaline (Greek, steno = narrow and haline = salt), which means they are restricted to either salt or fresh water and cannot survive in water with a different salt concentration than to that which they are adapted. However, there are exceptions- like the bull shark mentioned just a second ago.

Some salmon species are born in freshwater, then move to saltwater for several years, and then return to freshwater to spawn. Other fish, like some eels, are born in a marine environment, move to freshwater, and then go back out to sea to reproduce. If freshwater and saltwater fish kidneys work opposite of one another, how can there be fish that can do both?

Salmon returning upstream to spawn have many obstacles
to overcome. Their spawning grounds are usually a thousand
feet or more above sea level so they must leap up many
waterfalls. Oh, there are hungry bears too.
Salmon are famous for migrating to and from the sea. Almost all the species are semelparous (in Latin, semel = once and parous = breeding); this means that they return to their freshwater streams to spawn only once, and the trip and the reproduction kills them. The one exception is the Atlantic Salmon (Salmo salar). This species is spawned in, and returns to, the calm streams along the Atlantic coast several times in its life to spawn. This reproductive strategy is call iteroparity (itero = repeated). Iteroparous species lay fewer eggs at a time, the advantage is that survival chance is increased by repeated spawning – one bad year doesn’t destroy a big proportion of the population.

The migratory species of salmon are osmoregulators, as are most freshwater fish; their physiology demands a certain salinity level, and use energy to produce that level in their tissues. However, they can also adapt to various salinity levels. As such, these salmon as well as bull sharks are known as euryhaline (eu = good, haline = salt). Their physiology changes with the salt concentration.

While in freshwater the salmon will not drink, and will produce copious amounts of urine to get rid of the excess water it absorbs through osmosis.  But when it migrates to the ocean, it drinks all the time, and its kidneys work hard to remove the excess salts.

Chloride cells in euryhaline fish can sequester or
excrete salt, based on the hormone signals they receive.
This helps some fish move from aquatic to marine
environments and back again.
But the gills are the key to survival in the both the freshwater and saltwater environments. Energy consuming reactions will transport both Na+ and Cl- against their gradients, so they pump Na+ and Cl- into the fish’s tissues in freshwater and out of the fish’s tissues in saltwater. It is an adaptation of the marine fish’s chloride cells to work in both directions. This switch, as well as the kidney’s change in urine concentration, takes time. Therefore, salmon will spend days or weeks in intermediate zones, or estuaries, before going out to the ocean, and before returning to the rivers.

These are difficult lifestyle choices for salmon, the trips and the spawning kills them. So what is the advantage? The movement to oceans provides the growing salmon with readily available sources of food, so competition is reduced. The return to where they were spawned is just a good bet; if the stream was good enough to spawn them, then it is still probably a good place to lay eggs. Finally, working so hard to get to the spawning ground just a single time allows for selection of strong individuals, allows for huge numbers of eggs to be laid (the chance that some survive goes up), and the death and decomposition of the adults provides nutrients for the hatched fry (baby salmon). But these are human interpretations, I bet there are other advantages and disadvantages. However,  one thing is for sure, the balance sheet for these species comes out in favor of these adaptations – if it did not, nature would adapt further.

The eggs that don’t hatch and the carcasses of the mated
Adults create nutrient rich waters for the fry to develop in
before they head out to sea.
How about one more exception for today? Some individuals in semelparous species of salmon (Chinook, Coho, Pink, Steelhead, etc.) will not die after spawning, and will return again to the ocean. These individuals are often females, and are often smaller than average. These gals reverse their salt and water conservation strategies several times in their lives, making them prize winners for osmoregulatory exceptionality.

Next week, let’s tackle how the properties of hard water affect all life on Earth.

Sakamoto T, Ogawa S, Nishiyama Y, Akada C, Takahashi H, Watanabe T, Minakata H, & Sakamoto H (2015). Osmotic/ionic status of body fluids in the euryhaline cephalopod suggest possible parallel evolution of osmoregulation. Scientific reports, 5 PMID: 26403952

Cozzi RR, Robertson GN, Spieker M, Claus LN, Zaparilla GM, Garrow KL, & Marshall WS (2015). Paracellular pathway remodeling enhances sodium secretion by teleost fish in hypersaline environments. The Journal of experimental biology, 218 (Pt 8), 1259-69 PMID: 25750413

For more information and classroom activities on osmoregulation in fish and sharks, chloride cells, and reproduction strategies, see:

Osmoregulation in fish –

Chloride cells –

Osmoregulation in sharks –

semelparity and iteroparity –


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