Wednesday, August 31, 2016

Don’t Eat The Yellow Snow

Biology concepts – osmoregulation, tonicity, phytohormones, avian kidney, pinnieds, cetaceans

African elephants are larger than asian elephants, but their
urine production is similar. A 2007 study found that
African elephants can differentiate family members
based on their urine. It is similar to marking territory,
but they use urine to keep track of family members who
may be out of sight.
The asian elephant can urinate as much as 55 liters/day. That's about 3/4 of the volume of the average size bathtub! By comparison, the vaunted racehorse can only manage about 6 liters/day. Maybe we should rethink that old saying.

We know from the posts of the last few weeks that both salts and water are necessary for life, and that they work together to keep their concentrations within safe limits; a process called osmoregulation. You suspect correctly that kidneys and urination is involved, but what about plants – they don’t use the restroom.

For many animals, the kidney is the major organ of osmoregulation. The average adult human voids 1-2 liters of urine each day, but an uncontrolled diabetic with polyuria (poly=much and uria=urine) might expel 5-6 liters. Maybe we should bet on diabetics at the racetrack.

We get rid of water and soluble wastes via our kidneys. The kidneys filter the blood; nearly 800 liters of the red stuff each day. The basic filtering unit of the kidney is the nephron, who we met previously (Sorry, I Don’t Drink), made up of the Bowman’s capsule and sets of tubules.

Solutions of different tonicity have similar effects on plant
and animal cells, but plant cells can handle it better because
they have a rigid cell wall.
If the body is low on water, more water is reabsorbed in the tubules. Likewise, if the body has too much salt, few of the salt ions are reabsorbed in the tubules. In this way, our kidneys are basically concentrating our wastes in a small amount of water for excretion from the body. The amount of water depends on many factors, including the need to keep the cells at the right level of tonicity (concentration of salt relative to outside the cells).

Solutions can be hypertonic, meaning they have more salt than the cytoplasm, and water will flow out of cells by osmosis. Solutions can also be hypotonic, with less salt than in the cells (water will flow in to the cells) or isotonic, with the same osmotic pressure inside the cells as outside.

We all know that we don’t urinate the same amount all the time – drink more, go more. However, you don’t urinate the same amount you drink; your urine is concentrated by your kidneys in order to conserve water. Therefore, there must be some control mechanism. The answer is hormones. A hormone (“to set in motion” in Greek) is a small protein that is released from one cell and then acts as a chemical signal on other cells, either through the bloodstream (endocrine hormones) or through a duct (exocrine hormones) to the bloodstream or directly to other cells.

The angiotensin system. 1. The body senses that water
is low. 2. The kidney releases renin. 3. Renin  and
angiotensin converting enzyme produce angiotensin II
from angiotensin I in the lung. 4. Angiotensin II stimulates
aldosterone in the adrenal glands. 5. Aldosterone causes
more water and salt to be reabsorbed in the Loop of Henle;
this increases the blood volume and solves the problem.
Aldosterone is produced by the adrenal glands and acts on the distal collecting tubules of the kidneys. This endocrine hormone acts to conserve sodium and water and secrete postassium, thereby reducing urine volume but increasing the loss of potassium.  Aldosterone is released in response to angiotensin levels in the plasma, which in turn are controlled by sodium and water levels in the blood.

Arginine vasopressin (AVP, also called antidiuretic hormone) is another endocrine hormone that reduces the amount of water to be lost in the urine. This hormone is produced in the pituitary gland of the brain and also works to conserve water. By reducing the amount of water lost, the blood volume (which is mostly water) is increased, so blood pressure increases. This is why people are given intravenous fluids when they have lost a lot of blood.

The exceptions to this mechanism of kidney function are the mammals that live in hypertonic (saltwater) environments, like whales and dolphins (cetaceans) and seals or walruses (pinnipeds, latin for feather- or fin-footed). It is hard to study the urination in these animals in their native environment; they urinate in the ocean. Are you going to measure their individual contribution to the ocean – I think not.

Water wants to flow out of the cells and into the sea (hypertonic as compared to the cells), trying to balance the salt concentrations in both places. Therefore, the marine mammals must conserve freshwater or they become dehydrated. Both pinnipeds and cetaceans have large kidneys with enough renal tubule length to produce very concentrated urine, thereby conserving water. However, it appears cetaceans don’t really take advantage of this. Instead, they make a lot of metabolic water (Gimme Some Dihydromonoxide) and can keep from dehydrating by using the water they produce through cellular respiration.

Here is an inside view of a seal kidney. It’s huge! The many
lobules provides much tubular area to take up freshwater
and concentrate the urine.
Pinnipeds don’t drink water saltwater to any degree at all, they get their freshwater from their diet and their metabolic water.  Scientists use to think this was also true for cetaceans, but recent studies show that they do drink a small bit of seawater – not enough meet their water needs, but also not more than their kidney’s can handle.

Don’t think that marine (saltwater) mammals have it so bad. If they were to abandon the seas for freshwater sources, they would just trade one problem for another. Freshwater mammals have too much of a good thing, they run the risk of losing too much salt by being in so much salt poor (hypotonic) water all the time. This is why the kidney is so amazing, it can adapt functionally and anatomically to get rid of too much water or too much salt, depending on where you are. That is not to say the kidney is the only anatomic mechanism needed to maintain osmolarity within a tight range. Many organisms need more than kidneys, and have developed completely different mechanisms of osmoregulation.

Bird kidneys may be small, but they represent an evolutionary
intermediate, Some parts have short loops, like most mammals,
and some have long loops, like pinnipeds and cetaceans. However,
most of the kidney has reptile-like nephrons with long loops.
Birds share some water conserving and salt regulating apparatus with mammals. Avian (bird) kidneys have about 75% of their nephrons with reptilian structure, and 25% mammalian nephrons, containing a Loop of Henle. Therefore, avian kidneys are not as good at removing water and regulating salts as mammals are. Mammal urine can be concentrated 20-50x as compared to blood (the Kangaroo rat can produce a 9000x concentration), but birds can only manifest about a 2-3 fold concentration.

Therefore, birds have another mechanism to get rid of salt and maintain an osmotic potential within its limits. The salt gland is found in birds and reptiles. In many birds it is located near the eyes or nostrils (in crocodiles, salt is excreted through their tongues – everything tastes salty to them).  The salt gland removes Na+ and K+ from the blood, allowing birds and reptiles to consume saltwater or animals that live in saltwater.

Some organisms have it easier, like amphibians. With semi-permeable skin, they just leak salt out through their entire skin surface. Other organisms aren’t so lucky, like plants.

Plants must also regulate salt concentration, but they don’t have a familiar excretory system; in fact, they don’t have a specific osmoregulatory system. Water is lost via transpiration (Sorry, I Don’t Drink), and adjustments can be made to alter the amount of evaporation that occurs. Unfortunately, transpiration of water is linked to moving nutrients such as salts up the plant from the roots to the leaves. Therefore, shutting down transpiration will also shut down movement of nutrients. 

Plants in high temperature, low humidity, high wind environments have the highest rates of transpiration and are in danger of losing too much water. Once again, hormones are the answer. Plants do have hormones (phytohormones), so they probably have to deal with teenager issues just like human parents. Abscisic acid is an important hormone which shuts off transpiration. This phytohormone closes the stomata (stoma = mouth in Greek) on the upper sides of leaves, from which water evaporates and gases are exchanged. Abscisic acid also promotes water absorption from roots and root growth.

Some plants are cryptophytes by surviving unfavorable
seasons either underground (geophytes), hide their
seeds in the marshy mud (helophytes) or underwater
(hydrophytes). Hydrophytes in general are plants that
have their roots in water or water-logged soil.
Many xerophytes (plants that live in hot, dry places) have adapted to resolve these issues. They have leaf modifications to reduce water loss; needle-shaped leaves, sunken stomata, and waxy cuticles to cover the leaves. On the other hand, in hydrophytes (plants that live completely or almost completely in water), salts and water can be absorbed in the entire plant, not just the roots.

In terms of cations (Na+, K+), plants have a problem. They use potassium as their primary intracellular cation, but dirt is usually potassium-poor. Therefore, plants have K+ transporters to actively take up this ion. Unfortunately, the transporters don’t discriminate very well between K+ and Na+, so often times too much Na+ is taken up into plants.

Red mangroves have impermeable roots that help keep
out salt, and can also secrete some salt from there leaves,
but their most visible mechanism is the yellow salt leaves.

Excess Na+ can be toxic to cells, so measures must be taken to deal with these ions. Glycophytes are plants that are salt-sensitive, and include many of the plants that we cultivate. Therefore, soil salinity is an important factor in agriculture and gardening. Much research and breeding continues to an effort to produce crops that are better at differentiating uptake of K+ and Na+. Halophytes (halo=salt, phyte=loving), on the other hand, will allow the uptake of the excess ions, and then sequester them in vacuoles to prevent cellular damage.

Some plants live in extremely high salt environments. One example, the red mangrove tree, is a facultative halophyte. Facultative is a fancy way of saying “optionally.” These trees live in estuaries, where the river meets the sea. The water is quite salty there, and the mangroves are rooted in the water, so excess salt could be a problem. To deal with the toxicity of the excess Na+, the mangrove will store the salts in selected leaves, called the “kidney leaves.” When a toxic level is reached, the leaves turn yellow and just drop off. The tree must constantly invest energy in producing new leaves, so there is a cost to this way of life, but it seems to work for them.

If plants that live in or near seawater have adaptive mechanisms to maintain proper salt concentrations, then how about fish? We'll look at the osmoregulatory tricks by these organisms next week.

Ben Hamed-Laouti I, Arbelet-Bonnin D, De Bont L, Biligui B, Gakière B, Abdelly C, Ben Hamed K, & Bouteau F (2016). Comparison of NaCl-induced programmed cell death in the obligate halophyte Cakile maritima and the glycophyte Arabidospis thaliana. Plant science : an international journal of experimental plant biology, 247, 49-59 PMID: 27095399

Peña-Villalobos I, Valdés-Ferranty F, & Sabat P (2013). Osmoregulatory and metabolic costs of salt excretion in the Rufous-collared sparrow Zonotrichia capensis. Comparative biochemistry and physiology. Part A, Molecular & integrative physiology, 164 (2), 314-8 PMID: 23103672

Takei Y (2015). From aquatic to terrestrial life: evolution of the mechanisms for water acquisition. Zoological science, 32 (1), 1-7 PMID: 25660690

For more information, classroom activities or laboratories about osmoregulation, tonicity, abscisic acid, avian kidney, pinnipeds, cetaceans, see:

Osmoregulation –

tonicity and osmotic pressure –

abscisic acid –

avian kidney –

pinnipeds –

cetaceans –