Wednesday, March 18, 2015

The Search For The Unicorn - Slightly Off Center

Biology concepts – teeth, narwhals, unicorns, bilateral symmetry, evolution, mechanosensing, asymmetry



The movie Legend starred Tom Cruise and Mia Sara,
as well as a bunch of little people – you know, actors
that were small, not small actors. The unicorn pair
represented light and goodness, and kept the devil
at bay. Until Mia got cocky and touched one. Then
Cruise had to save the day.
It’s no secret that some pretty odd and awful stories have come out of North Korea in the past few years. Kim Jung Un and his recent ancestors have done some amazing things….. supposedly. Un’s father, Kim Jung Il apparently invented the hamburger, and he shot 11 hole-in-ones in his first round of golf.

Not to be outdone, Kim Jung Un made an amazing announcement in 2012. He and his archeologists discovered a unicorn lair. Yep, North Korea’s twenty-something leader proved the existence of unicorns. The lair was supposedly the resting place of the unicorn ridden by the great King Dongmyeong, around the year 0 CE.

The earliest writings that describe unicorns were those of the Greek, Ctesias, in the late 5th century BCE. He described the Indian Ass, an animal with a white, strong body and perhaps a red head from which sprung a long single horn of red, white, and black. It was said that a cup made from the horn could neutralize any poison.


There are real animals with one horn, like the
unicorn leatherjacket fish in the top left, and the
Indian rhinoceros at the bottom left. The rhinoceros
beetle has one big horn and fairly large part of his
jaw below, so I don’t know if he counts. On the top
right is the Meller’s chameleon. They say he a has a
horn on his nose, but you have to look close and
want to see it.
Four hundred and fifty years later, Pliny the Elder, historian of Rome, also wrote about a very strong animal with a single horn protruding from its forehead. He described an oryx (an antelope with a single horn), an Indian Ox (probably a rhinoceros – rhino = nose and ceros = horn), and the same Indian Ass with a horse-like build and a single horn.

Pliny wrote, “The unicorn (uni = one, and ceros = horn) is the fiercest animal, and it is said that it is impossible to capture one alive. It has the body of a horse, the head of a stag, the feet of an elephant, the tail of a boar, and a single black horn three feet long in the middle of its forehead. Its cry is a deep bellow.” Uh-huh. That doesn’t sound much like an antelope or a rhino, so I guess he meant the Indian Ass.

Soon, Romans were trading long spiral tusks, but no one was telling where exactly they had come from. These “unicorn” horns were snow white with a tight spiral. As a result of these horns, the unicorn in the West settled down to be a pure white horse with a very long, pure white, spiraled horn. This is the image we generally see in tapestries and illustrations.


Kirin Beer from Japan uses a unicorn (kirin) as its
logo. Look closely and you can see the single
horn on its head.
In the Far East there were unicorns as well. Known as the qilin (pronounced chee-lin) in China, there was a version in Japan too, the kirin. This was a benevolent animal, with shiny scales like a dragon and one or perhaps two horns. It avoided fighting and walked so softly that it would not disturb or harm a blade of grass. An animal like this (perhaps the saola) is most likely the one referred to in the North Korea stories.

But what about real life? Most likely, those horns in the Roman markets were really narwhal tusks, as discussed in a 2011 paper. It is very likely that the narwhal played into the unicorn legend, as their tusks could be offered as concrete proof of unicorn existence.

The narwhal (Monodon monoceros) is an amazing animal, and fits into our recent theme of animals that abandon bilateral symmetry. Monodon means one tooth, and monoceros means one horn; a pretty accurate name, all in all.


Our post today uncovers many of the problems
with these cartoon narwhals. Yes, they love where
there is ice. But they don’t have all those teeth, the
tusk isn’t centered and doesn’t come out of their
forehead, and they don’t have a dorsal fin
to speak of.
Narwhals are a species of whale, meaning that they are mammals. They live way up north. From Baffin Bay, around Greenland, to the north of Russian, they swim in pods of 10-100, but you’ll rarely see them even if you live near there. There are perhaps 45,000-50,000 narwhals today.

This is a steady number because it’s so hard to get to where they live. Consequently, narwhals haven’t been hunted into extinction. They spend a lot of their time on deep dives under the ice floes, so they aren’t seen often. No narwhal has ever been seen feeding; we only know what they eat from examining stomach contents.

Their most distinctive feature is the long (up to 10 ft/3 m) tusk on the males. Just one tusk, mind you, like a unicorn horn. The narwhal tusk - like elephant, walrus or warthog tusks - is a tooth.

Very young narwhals have six maxillary (upper jaw) tooth buds and two pairs of tooth buds in the lower jaw (mandible). However, only one pair develops any further. A tooth bud is what you find on an X-ray of a child (see picture).


You can see the teeth developing from crown to
root in the darker tooth buds. The pulp is usually
dark, but the middle tooth has had a root canal
and a filling has been placed in the whole pulp
chamber. The large tooth to eh left is the first
molar. It doesn’t have a baby tooth to push out
of its way.
Teeth form in the jawbones as tooth buds. Most narwhal teeth never go past the tooth bud stage, but occasionally a tooth will erupt where one shouldn’t. These are often misshapen or caught between the bone and the palate, or in the wrong place. This is all good evidence that the teeth are vestigial; they serve no functional purpose for the normal narwhal.

Just one tooth, almost always the left cuspid (most people call it a canine), does develop. Hold on though, it isn’t that simple. Instead of developing in a vertically directed tooth bud and erupting down through the jaw, the left canine stays horizontal and erupt right through the front of the jaw and through the narwhals lip!

Since the tusk is derived from the left cuspid, it erupts left of center, making the narwhal bilaterally asymmetric! A 2012 study showed that the bony attachment and length proves that the narwhal tusk is a canine, not an incisor as so many people think. But, it’s not just the location that makes the narwhal tusk amazing, it’s how it’s made and what it can do.

A 1988 study suggests that the tight spiral as it grows keep the tusk from curving. A curved tusk would make it hard of the narwhal to swim in a straight line. Whatever the reason, the spiral is an iconic image for both narwhals and unicorns.


The top image shows how the narwhal tusk is off
center. The bottom image is my analogy. The tusk
is offset like a knight with his jousting lance. This
is Heath Ledger in A Knight’s Tale. Um….why isn’t
he wearing armor?
Despite being a tooth, the tusk is quite flexible. It can bend up to a foot (0.3 m) in any direction without breaking. It’s awfully long, we said 10 ft. above, but most are in the 8-9 foot range. This is huge when you think that most male narwhals are only about 15 foot long in the body.

Teeth are normally built with the hard enamel on the outside. Enamel is harder than bone and protects the teeth from breakage when chewing. The mouth is a rough environment and teeth have to put up with a lot of abuse.

Deep to the enamel is a material called dentin. This stuff has a lot of similarity to bone, although it isn’t quite as hard and doesn’t have living cells within it (like osteocytes – see this post). The dentin does contain millions of tubules that go from the enamel junction all the way to the pulp in the center. The pulp has a nerve and blood vessels.

The dentinal tubules have fluid and small processes of the neuron in them. When you eat something cold or have a cavity, the fluid in these tubules moves and changes the pressure in the pulp chamber. The single neuron in the tooth is a pain neuron, so any pressure change is interpreted by your brain as pain. It teaches you to take care of your teeth, but it ain’t the most pleasant of all evolutionary adaptations.


The cartoon on the left shows the enamel crown
covering the dentin and the dentinal tubules.
Inside the tubules are the odontoblasts that lay
down dentin all during the life of the tooth and the
nerves that go into the tubules. The right image is
an electron photomicrograph of the tubules.
The narwhal tusk is different. It is the only tooth known that has the dentin on the outside, although a 1987 study showed that it has no enamel, so it isn’t really an inside out tooth. The dentin is covered by a thin layer of cementum. This is what normally covers the roots of the teeth and helps attach them to bone. The dentin of the narwhal tusk has about 10 million of those tubules, but it is different from human dentin.

A 1990 study compared calcium content and hardness between human teeth and narwhals. The narwhal cementum was more mineralized than human, but the dentin of narwhals was less mineralized than human dentin and was softer. This may be why the narwhal tusk is so flexible.

The tubules of the narwhal tusk dentin connect to channels in the cementum, so there is a communication to the outside. A group in 2014 showed this and used the information to hypothesize that the tusk is a mechanosensor. Experiments showed that their heart rate changed when the water touching the tusk was switched from freshwater to salt water. They hypothesize that the tusk senses temperature, salinity, pressure, and perhaps touch to help in navigation and hunting.

But if that’s the case, why do only males have them? Females have to hunt too. The group from the 2014 paper offers that males and females have sexually dimorphic foraging techniques – they eat different things and hunt differently, so females don’t need horns. This is not well-supported. Many scientists believe the long tusk is a sign of health and genes and is therefore an ornament for mate selection.


The dorsal fin of the narwhal is greatly reduced. It
has notches that scientists hop to use to identify
individuals. The lack of a dorsal fin is believed to
be so they don’t injure it on the under side of the
ice floes when they surface, but it could also be so
they don’t run it into the ocean floor as they feed
upside down.
Occasionally, one will see females with a tusk, but like with many tusked females (elephants, etc), they are usually shorter. You can also find narwhal males with two tusks. But two tusks doesn’t mean that they are returned to bilateral symmetry. Both tusks spiral to the left! There must be some strong left-hand genes at work.

One last thing. The offset tusk lead to another weird narwhal behavior. A group in 2007 put cameras and positional monitors on some narwhals and found that they tend to swim upside down a lot. Almost 70% of their time on the ocean floor was spent in the supine position. Since the tusk points down just slightly, scientists believe they hunt upside down so that the tusk won’t get stuck in the ocean floor and break! The tusk must be pretty important - or they just like lounging on their backs.

Next week – another whale has become asymmetric, but in a completely different way. This time, it’s the nose that goes.



Christen AG, & Christen JA (2011). The unicorn and the narwhal: a tale of the tooth. Journal of the history of dentistry, 59 (3), 135-42 PMID: 22372187

Kingsley, M., & Ramsay, M. (1988). The Spiral in the Tusk of the Narwhal ARCTIC, 41 (3) DOI: 10.14430/arctic1723

Nweeia, M., Eichmiller, F., Hauschka, P., Donahue, G., Orr, J., Ferguson, S., Watt, C., Mead, J., Potter, C., Dietz, R., Giuseppetti, A., Black, S., Trachtenberg, A., & Kuo, W. (2014). Sensory ability in the narwhal tooth organ system The Anatomical Record, 297 (4), 599-617 DOI: 10.1002/ar.22886

Dietz, R., Shapiro, A., Bakhtiari, M., Orr, J., Tyack, P., Richard, P., Eskesen, I., & Marshall, G. (2007). Upside-down swimming behaviour of free-ranging narwhals BMC Ecology, 7 (1) DOI: 10.1186/1472-6785-7-14




For more information or classroom activities, see:

Narwhals –

Tooth structure –



Wednesday, March 11, 2015

The Eyes Have It

Biology concepts – asymmetry, lateral polymorphism, flatfish, evolution, copepod, ecology, niche



Ray Harryhausen was the most famous of the stop
motion artists in the movies. This version of the
Cyclops was his creation for the 1958 movie, The 7th
Voyage of Sinbad. I can’t see how the Cyclops could
catch anything with just the one eye – he had no
depth perception.
We have been talking about bilateral symmetry in the past few weeks, and this would include having two eyes, one on each half of your face. Two eyes must be a pretty important evolutionary adaptation; can you think of an animal that has just one eye – other than a cyclops, that is? (the answer is somewhere in post) Some protists have a single eyespot for sensing light – but they aren’t animals.

Predators need to catch their food, so they need depth perception. For this you need two overlapping images. You don’t do the math consciously, but your brain uses the differences in each image to tell you how far away the target prey is. To get two images simultaneously, you need both eyes to be on the front of your face.

Prey animals, usually herbivores, have to worry about being chased down by a predator. Prey animals are usually quick, but they need clues to get a good start before the predator gets too close. Their eyes are designed to pick up motion; it doesn’t matter how far away it is. If something moves in their line of sight, they assume it’s a predator and they bolt. By having their eyes on the sides of their head, they have a maximal range of vision which gives them the best chance to see that lioness coming.

Just to show you how important evolution thinks it is to have at least two eyes, let’s discuss a group of animals that have found a way to make two drastic changes work for them. Flatfish have evolved to lie on their side, but they’re predators so they can’t afford to have one eye constantly seeing nothing but the sandy ocean bottom. Consequently, they have moved one eye to the other side of their head!


This is a winter flounder (Pseudopleuronectes
americanus). Notice how it blends in to the floor. It
does more by flapping and tossing sand on its back.
The mouth points up when the fish is vertical, so the
left most eye is the one that migrated. Makes sense,
you wouldn’t want to move an eye under your chin –
that would be silly.
Flatfish are all from the order Pleuronectiformes (pleuro = toward the side), sometimes called the Heterosomata (hetero = differently, and soma = bodied). There are some 715 species of flatfish in 11 families, and they include the turbots, sole, flounder, plaice, and halibut. They are all predators that lie in wait on the ocean-, lake-, sea-, or riverbed. But they don’t start out that way.

All the Pleuronectifromes start out as fry that swim upright. Their top is at the top, and the bottom is toward the bottom. They live nearer the top of the water than the bottom, have pigment coloration on both sides of their body, and feed on phyto- and zooplankton. But the surface isn’t the safest place to live; bigger fish are always around to eat the small fry.

So, as they start to develop into adults, many changes take place. They swim down to the floor of whatever body of water they call home. One eye starts to move! It travels over the top of their head and onto the other side, like the Mr. Potato Head of some deranged child.

As you can imagine, moving an eye isn’t an easy thing to do. Their brain has to move, as do the cranial and facial bones. Their mouth has to make room for the eye coming its way. All in all, it’s a tough piece of work.


The top image is the visible side of a rock sole, the
bottom image is the lower side of the same fish. As the
fish matures and lies on its side, the coloring changes.
Pigments are energetically expensive to make, so why
waste them. The middle image shows the line where it
goes from pigmented to unpigmented. What control!
Moving an eye from one side of the head to the other seems illogical, why not just develop with both eyes on one side, or make do with one eye? Creationists have used the flatfishes as an argument against evolution. We have fish with eyes on each side of their head, and fish with both eyes on one side of their head. They argued that if natural selection was responsible for the change in flatfish, there should be fossils or fish that are intermediates.

Well, there are evolutionary intermediates – even some living examples. A paper in 2008 introduced us to two extinct species of flatfish. In each (Amphisitium and Heteronectes), one eye had migrated, making the fish asymmetric, but it had not crossed the crown of the skull and made it to the other side. These are definitely intermediate species, but we can go one better.

The Psettodes genus of flatfish (the turbots - yummy by the way) have one eye that is located right at the crown of their skull. Perhaps technically you could say that is has migrated to the other side, but just barely. And this brings us to another question, which eye moves?

Some flatfish are right-eyed (dextral) that swim left side down. Others are left-eyed (sinistral) and swim with their right side down. Some species are strict and some are more likely to have reversants (individuals that lay on their other side). A 2005 paper stated that only about 7 of the 700 species of flatfish show lateral polymorphism (lateral = side, poly = more than one, and morph = shape), ie. some right-eyed and some left-eyed individuals.


The bottom image has on the left side depicts the two
extinct species of flatfish where the migrating eye hadn’t
quite made it to its destination. The middle cartoon is
the extant Psettodidae, like the Indian Halibut on the top.
The eye has just made it to the crest of the cranium. The
right cartoon on bottom shows a species where the eye
has completely migrated. Looks like great support
for evolution.
For example, a 2009 paper describes reversants for two right-eyed flounder species which are the first left-eyed individuals ever seen in these two species (Microstomus achne and Cleisthenes pinetorum). It goes the other way too – a 2013 study describes the first right-eyed individual ever seen in a megrim (sometimes called a whiff, Lepidorhombus whiffiagonis). But in the left-eyed California halibut (Paralicythys californicus), up to 40% of the individuals are right-eyed. Perhaps right-eyed species are stricter than left-eyed species.

The exception to that rule is the starry flounder (Platichthys stellatus). It’s a member of a right-eyed family of flounders, but in some cases half of the individuals are left-eyed. In this case, there seems to be more to the story - the lateral polymorphism occurs only in populations of specific geographic areas.

A paper from 2007 looked into the mystery that 7 species that show lateral polymorphism, but only two (starry flounder and P. fleusus) show a geographical distribution in their polymorphs. Off the coasts of Japan and Russian, 100% of the starry flounders are left-eyed, but near Alaska they are only 75% sinistral and from Washington state to central California the populations are about 50/50.

The researchers looked into several questions. Was there more to being left-eyed or right eyed than just which side the eye went to? Does the side make a difference that could account for the geographic distribution? They found out – yes, and yes.

They saw that the right and left-eyed individuals have more asymmetries between them than just the side of the body that the eye migrates to. They have differences in mouth size and angle, as well as tail size of all things. Right-eyed individuals had significantly longer and wider tails than did left-eyed individuals!

The research also shows that in areas where the two groups compete the most, the differences between the dextral and sinistral individuals are the greatest. This suggested to them that the differences allow them to compete for different prey – to fill different ecological niches. The hypothesis is that the polymorphic asymmetries give them different advantages which they then exploit and this is why there is a stable geographic distribution in populations.


A very complicated but informative chart from the paper
on pitx2 reactivation during metamorphosis. Note which
side is right and left and then follow what happens for
the two species, one right-eyed and one left-eyed. The
right side shows what can happen if you block pitx2. In
the reversed individual, the left habenula (dd. l) enlarges
instead of the right.
Well, that’s cool, but how is it controlled – what makes a fish right- or left-eyed? A 2009 paper started to investigate this. During development of the embryo, certain internal asymmetries develop (will talk more about this in a few weeks). In fish and other vertebrates, this is controlled by expression of certain genes in the habenula of the brain. One of the gene products (proteins) in the habenula is called pitx2.

There are actually two habenulae, one in each hemisphere of the brain, as part of the thalamus. The researchers looked at the brains of right-eyed and left-eyed flounder species and detected some things that were the same and some things that were different.

After the pitx2 did its job in the embryo, it was turned off. But right before metamorphosis – when the eye migrates and the fish lies down on one side, pitx2 was turned back on – only in the left habenula, regardless of which way the fish’s eye was going to migrate and which way it was going to lie down.

Not only that, but the right habenula started to grow bigger than the left habenula in both dextral and sinistral species. The only thing that was different was the rotation of the brain, with the left habenula moving forward in the right-eyed species and the opposite occurring in the left-eyed species. The same changes occurred in fish no matter which eye migrated!


This copepod is about to swim into the top border of
this image. The reason for this is that he only has one eye
(the light spot between the antennae), so he has no depth
perception! By the way, he’s only 2 mm long, so he
probably will just bounce off the edge.
However, in those fish that they experimentally blocked the second round of pitx2 activity, you could get a normally turned fish or a reversant. The only difference - when you got a reversant, it was the left habenula that enlarged, not the right. I think we still have some more to learn.

By the way – at the beginning of today’s post I asked if there were any real animals with one eye. I made it sound like there aren’t but in fact there are exceptions. In some small crustaceans (arthropods) called copepods, the majority of species have a single eye right in the middle of their head! What’s more, a 1994 study showed that the holdfast of a particular copepod parasite is asymmetric (like we saw last week) and this particular copepod is a parasite of only flatfishes! How symmetric this tale of asymmetry turned out to be!

Next week - can a single tooth render an animal asymmetric? Well, that depends on the tooth, doesn't it.



MacDonald P (2013). A rare occurrence of reversal in the common megrim Lepidorhombus whiffiagonis (Pleuronectiformes: Scophthalmidae) in the northern North Sea. Journal of fish biology, 83 (3), 691-4 PMID: 23991885

Suzuki, T., Washio, Y., Aritaki, M., Fujinami, Y., Shimizu, D., Uji, S., & Hashimoto, H. (2009). Metamorphic pitx2 expression in the left habenula correlated with lateralization of eye-sidedness in flounder Development, Growth & Differentiation, 51 (9), 797-808 DOI: 10.1111/j.1440-169X.2009.01139.x

Goto T (2009). Reversals in two dextral flounder species, Microstomus achne and Cleisthenes pinetorum (Pleuronectida; Teleostei), from Japan. Journal of fish biology, 74 (3), 669-73 PMID: 20735586

BERGSTROM, C. (2007). Morphological evidence of correlational selection and ecological segregation between dextral and sinistral forms in a polymorphic flatfish, Platichthys stellatus Journal of Evolutionary Biology, 20 (3), 1104-1114 DOI: 10.1111/j.1420-9101.2006.01290.x

Friedman, M. (2008). The evolutionary origin of flatfish asymmetry Nature, 454 (7201), 209-212 DOI: 10.1038/nature07108


For more information or classroom activities, see:

Flatfish –

Copepods –

Habenula -




Wednesday, March 4, 2015

Looking Sideways In The Mirror

Biology Concepts – platyhelminthes, asymmetry, bilateral symmetry, evolution, cephalization, natural selection, fish, lepidophagy

What is the largest living structure on Earth? No, it’s not the 2200 acre Armillaria ostoyae fungus in Oregon that we talked about previously. That is the largest single organism, but there is something bigger.

The Great Barrier Reef houses more species of
coral than any other place on earth, more than 600
species call the reef home. You see how many shapes
they can take. Does this mean they are
asymmetric animals?
The Great Barrier Reef off the northeast coast of Australia is alive. Reefs are made of the exoskeletons of coral polyps, with the new corals growing next to and on top of the older ones. With all the nooks and crannies available, coral reefs are some of the most diverse ecosystems on Earth, with thousands of species per square mile.

This can add up quickly, because the Great Barrier Reef is more than 132,973 sq. miles (344,400 sq. km) in area. And if you still don’t believe it is a living structure, get this; it’s moving south! Climate change is warming the waters off the coast, and corals and tropical fish are moving south with the warmer temperature. Meanwhile, the northern edge recedes as the waters get too hot for corals.

Corals take all sorts of shapes (see picture above), and despite what we talked about last week, they don’t seem to be either bilaterally or radial symmetric. Sure, brain corals look radial, but most corals don’t have a repetitive shape. Are these asymmetric animals?

Nope - remember that the coral you see is the exoskeleton of the polyp, not the animal itself. Just because the apartment building isn’t symmetric, it doesn’t mean the animal is as well. Coral polyps are definitely radially symmetric, so our discussion of last week still holds.

Corals and other radial animals are exceptions, since 99% of animals are actually bilaterally symmetric. But there are exceptions with the bilateral animals as well. Some species that have been bilateral for millions of years then evolved a tweak to the system. Some part them became asymmetric in order to give them an advantage. Their stories are exceptional and we should explore some of them.

Coral polyps are cnidarian animals. They live inside
the calcium shells they produce, and it is the shells
that seem asymmetric in many cases. But the polyps
show radial symmetry. Each stalk and white “flower”
is an individual polyp.
But we begin with a challenge –as we discuss the different animals that break symmetry in the next few posts, see if you can find something else that is common to many of them.

Let’s start with the first animals that became bilaterally symmetric– the platyhelminthes, or flatworms. Most flatworms are small, just barely visible with the human eye, and most swim in the water. There are free-living versions and parasitic species and it is in the parasites that we find our first animal that has decided that completely bilateral isn’t necessary.

Polyopsithocotylea monogenea is a group of flatworms that live on and feed on fish gills. About 0.05-1 mm long, these platyhelminthes attach themselves to one of the gill ridges and take up residence there for life. The attachment they use is called a haptor, and they come in different shapes and sizes.

The oncomiracidium stage of the worm is directly after the egg stage and before it becomes an adult. This stage is completely bilaterally symmetric. But when the adult stage is reached and it’s time to settle down and starting feeding on some fish’s gills, they become asymmetric via their haptor attachment.

Opisthaptors, or just haptors, are the attachment
organs for many parasites, including parasitic
flatworms. They can use suckers or clamps and hooks
in order to anchor the worm in its preferred habitat.
You can see that these haptors maintain animal
symmetry, but not all do.
Different host species fish have different gill anatomy, so the attachment point and position will be different. The haptor(s) have to be located where attachment is possible. This means that they may be on one side of the body or the other, or two on one side and one on the other, etc.

The initial haptor is usually located on the posterior end, on one side only. So much for bilateral symmetry. In some species this haptor has suckers, in others it has hard clamps, and yet other species have both.

As the adult grows, more haptors may develop, just where the animal touches the gill. Some may have 50 or more haptors arranged around their posterior, becoming more and more asymmetric. But there is a plan, they only grow where attachment is possible; some signal is generated by contact and this stimulates growth of more attachments.

Here we have a family of parasitic worms that aren’t symmetric living within a phylum of worms that were the first to be bilaterally symmetric - exceptional. But, take one step up the chain and you see the fish they live on. It just so happens that at least one group of fish parasitized by P. monogean worms are asymmetric themselves.

These are the profiles of some monogean flatworms. The
haptors of these parasites grown in odd places and
destroy the bilateral symmetry of the animal. But it is
necessary for the worm to attach to the gills of their
prey fish.
Cichlid fish are one of the most diverse family of animals known, with more than 1700 known species. They are found in the Old World and the New. They exhibit some amazing adaptations, especially when many are found in one place, Lake Tanganyika on the border of Tanzania and the Democratic Republic of the Congo.

The cichlids are successful because different species have developed different feeding niches, and this is where we meet our asymmetric cichlids – they eat the scales off other fish! One genus, Perissodus, has at least six species that eat scales, all endemic to Lake Tangayiki (although there are also other scale eaters in other locations).

Scale ripping and eating is called lepidophagy (lepido = scale, and phagy = eat). Scales are an unexpectedly good source of nutrition. They chock full of protein and calcium phosphate, and their attachments are both cartilaginous, fatty, and come with some carbohydrate. Remember that the next time you order fish in a fancy restaurant. Tell the chef not to scale it – you’ll be quite the topic of conversation in the kitchen.

Fish scales have many uses. Some, especially from
herrings, are used to make the pearlescent cosmetics
that are sold today. Gals, your putting fish scales on your
eyes and lips. Fish scales can also be turned into artificial
bone, or they can be food for lepidophageous fish.
Mind you, this isn’t eating the scales of dead fish, or eating the scales that drop off live fish. Lepidophagy means eating the scales that are still attached to live fish. It’s a fish smorgasbord. This is both good and bad. Scales on live fish grow back fast, so there is always a ready supply of food.

But, you can imagine that the fish being unfrocked don’t appreciate it very much and fight back or swim away quickly. That doesn’t even take into account how hard it is to bite the scales off a swimming fish. Therefore, lepidophages must evolve anatomies and behaviors that give them a chance to succeed. Or perhaps it would be better to say, they acquired characteristics that made being a lepidophage an advantage.

Here are two P. microlepis scale eating fish. One is right
mouthed and the other is left mouthed. You can see how
the way their mouth develops breaks their bilateral
symmetry. The right-mouthed version (on the left) will
only eat scales from the left side of fish. How is it that both
versions can be maintained in a population?
The species P. microlepis has developed one particularly amazing way to help it eat the scales off of neighbor fish. It’s mouth and jaws have evolved so that they bend sideways. There are right-mouthed P. microlepis and left-mouthed individuals. The difference is obvious, right-mouthed individuals will feed only from the left side of their prey and vice versa. They have an asymmetry, or lateralization, of both anatomy and behavior

A 2012 video study showed that right-mouthed individuals almost always attack prey from the left, and their strikes are more powerful and successful when coming from the preferred side. One could ask, why are their both types? How did right- and left-mouthed individuals come to evolve and why are there still both types?

A different 2012 study shows that juveniles prefer one side or the other, even before their mouth bend has become pronounced, so it is a deeply penetrating characteristic, both heritable and perhaps partially acquired. There’s evolution and genetics at work here.

The prevailing model is that at any one time, right or left-mouthed individuals will predominate in a population. Let’s say that right now, right-mouthed feeders are the majority. The prey fish will learn to pay more attention to their left side, as this side is more vulnerable.

This makes it harder for right-mouthed individuals to feed, but easier for left-mouthed fish, because the prey fish ignore their right side relative to the left. In time, the right-mouthed individuals breed less well and the numbers in the population will shift. The left-mouthed feeders will become the majority. This is an example of negative frequency-dependent selection, where as a trait becomes more common it becomes less advantageous, and there is a balancing selection.

The flu virus comes with one of many hemagglutin proteins
and one of many neuraminidase proteins. If one genetic
version is too successful, most people will develop
immunity to it and it becomes less fit in the population of
hosts. Its success is its downfall and a more rare version
will rise up. This is negative frequency-dependent selection.
The cycle will begin again after left-mouthed individuals come to dominant in the population. Back and forth the population will go. If the population stayed 50/50, nobody would gain an advantage, and overall, they would all suffer. The system only works if a small number develop opposite to the majority. The gene regulatory complex that controls if an individual will be right- or left-mouthed must be very complex if it can take into account that a few need to be lateralized the other way.

However, a 2012 study throws this into question. They found equal distributions of right-and left mouthed individuals in five populations they studied. Also, left-mouthed and right-mouthed individuals mated with each other just as often as they mate with same-sided individuals (called disassortative mating). Perhaps the mouth bend is not a true dimorphism (di = two, and morph = shape).

Or, as a 2008 study suggests, negative frequency-dependent selection works best when there is disassortative mating. This may be necessary since a 2007 study showed that lefty:lefty matings give 2:1 lefty offspring, right:lefty mates give equal righty and lefty offspring, but righty:righty pairs ONLY give righty kids. Figure out the genetics of that. The authors proposed two possibilities – mendelian genetics with lefty being dominant and dominant homozygous being lethal, or cross-incompatibility that is predominant in lefty:lefty homozygotes, (meaning lefty homozygotes can’t mate successfully mate with the other types).

Today we have seen a swimming flatworm that feeds on some fish, and some fish that feed on the scales of other fish. And both of them achieve this only because they have adapted their bilateral symmetry to become just a bit asymmetric.

Next week, there are other animals that break symmetry to survive. Flatfish lay on their sides at the bottom of lakes and oceans, yet they still use binocular vision. How can that be?



Takeuchi, Y., Hori, M., & Oda, Y. (2012). Lateralized Kinematics of Predation Behavior in a Lake Tanganyika Scale-Eating Cichlid Fish PLoS ONE, 7 (1) DOI: 10.1371/journal.pone.0029272

Lee, H., Kusche, H., & Meyer, A. (2012). Handed Foraging Behavior in Scale-Eating Cichlid Fish: Its Potential Role in Shaping Morphological Asymmetry PLoS ONE, 7 (9) DOI: 10.1371/journal.pone.0044670

Kusche, H., Lee, H., & Meyer, A. (2012). Mouth asymmetry in the textbook example of scale-eating cichlid fish is not a discrete dimorphism after all Proceedings of the Royal Society B: Biological Sciences, 279 (1748), 4715-4723 DOI: 10.1098/rspb.2012.2082

Takahashi, T., & Hori, M. (2008). Evidence of disassortative mating in a Tanganyikan cichlid fish and its role in the maintenance of intrapopulation dimorphism Biology Letters, 4 (5), 497-499 DOI: 10.1098/rsbl.2008.0244

Hori, M., Ochi, H., & Kohda, M. (2007). Inheritance Pattern of Lateral Dimorphism in Two Cichlids (a Scale Eater, Perissodus microlepis, and an Herbivore, Neolamprologus moorii) in Lake Tanganyika Zoological Science, 24 (5), 486-492 DOI: 10.2108/zsj.24.486



For more information or classroom activities, see:

Great Barrier Reef –

Coral polyps –

Cichlids –