Thoughts of a clam

To us active, dynamic mammals, the humble clam can appear positively…inanimate. Their nervous system is decentralized relative to ours, lacking any sort of brain, and to the untrained eye, it can appear that their only discernible reaction to the outside world is opening or closing. Open = happy, closed = not happy; end of story, right? Some vegans even argue that the clams are so nonsentient that it is okay to eat them and think of them as having no more agency than a vegetable!

You might already have predicted I intend to tell you about just how animate and sentient clams can be. But let’s start out by describing the nuts and bolts of their nervous system. As with many invertebrates, their nervous system is distributed throughout their body as a system of ganglia. Ganglia are clumps of nerve cells which may have local specialization, and transmit messages within neurons using electrical potentials. At the connection between cells (called a synapse), neurotransmitters are used to pass signals to the next cell. Researchers have found that bivalves use “histamine‐, octopamine‐, gamma‐aminobutyric acid‐ (GABA)…like immunoreactivity” in their central and peripheral nervous systems, much like us vertebrates do, and other studies have even found that the response to serotonin and dopamine is localized in nervous tissue linked to different organ systems.

Nerve cells (bright green) highlighted in a larval oyster with fluorescent dye (from Yurchenko et al 2018)

These systems of chemical nerve transmission are truly ancient, likely dating back to the formation of complex animal body plans in the earliest Cambrian. Researchers have great interest in studying these nervous and hormonal signaling systems in mollusks because they can shed light on the relative flexibility and limitations of these systems throughout the animal tree of life. Characterizing these systems can also allow us to understand the mechanisms that bivalves and other animals use to react to environmental stimuli.

Electron microscope view of gill cilia, zoomed in 1000x (from Dan Hornbach)

Like humans, bivalves spend a lot of time and effort eating. Most bivalves eat by filtering food from passing water with tiny cilia on their gills. These cilia work to capture food particles and also act as a miniature rowing team moving water along the gill surface. The bivalve needs a way to control this ciliar activity, and researchers found they could directly control the speed at which oysters move their cilia by dosing them with serotonin and dopamine, which respectively increased and decreased activity.

Bivalves also work very hard to make babies. Most bivalves reproduce by releasing sperm and eggs to fertilize externally in the water column. To maximize their chances to find a mate, they typically save up their reproductive cells in gonads for multiple months and release them in a coordinated mass spawning event. It appears that this process is controlled by hormonal releases of dopamine and serotonin. Researchers have determined that serotonin concentrations vary through the year, with mussels in New England using it to regulate a seasonal cycle of feeding in summer, followed storing of that energy for winter. During the winter when food is less available, they use that stored energy to bulk up their gonads in time for reproductive release in spring months, when their larvae have plentiful access to food and oxygen, ensuring them the best chance of survival. In recent decades, aquaculturists have learned to use serotonin injections to induce spawning in cultured clams, to ensure they will have a harvest ready at a certain time of year.

So bivalves are very sensitive to the seasons. How about shorter term sources of excitement? You might have observed this yourself through the clam’s most iconic activity: opening and closing its shell. Clams close their shells with powerful adductor muscles which pull the two valves together. A springy ligament at the hinge pulls the shell open when the muscles relax. Just like us, the clam needs to use nerve cells to signal the muscle to do its thing. In addition, two different sets of ganglia act to control the foot that some bivalves can extend to dig into sand, with one ganglion acting to extend the foot and the other causing it to contract. While clams don’t have a centralized brain with specialized regions for different uses like we have, this represents a sort of specialization of neural systems with a similar result.

Clam licking salt - Imgur
This iconic gif is often shared along with the claim it shows a clam “licking” salt. It is actually using its foot to search for a place to dig. The salt was not needed.

When a certain neuron is used repeatedly, it can form a cellular memory allowing the organism to acclamate (ugh sorry) and moderate its response to a particular stimulus over time. Giant clams, for example, close their shells when their simple eyes detect a shadow overhead. This behavior can protect them from predation. When I conducted some of my PhD research, sampling body fluid of aquarium and wild giant clams with a syringe, I noticed that captive clams didn’t close up in response to my shadow overhead, while wild clams required me to sneak up and wedge their shells open with a wooden block to do my work. I suspected that after exposure to frequent feedings and water changes by aquarists, the clam had “learned” that there was no reason to expend energy closing its shell. Meanwhile, in the process of proving that our sampling technique was not harmful to the animal, I discovered that clams which detected my shadow would quickly reopen within seconds when I hid from them, while those that were stuck by a syringe would stay closed for minutes before opening and beginning to feed again. Makes sense!

Other researchers noticed this phenomenon as well. One group found that giant clams repeatedly exposed to shadows of different sizes, shell tapping and even directly touching its soft tissue began to habituate (become accustomed) to the stress, opening more quickly and staying open longer each time the stimulus occurred. Even more interestingly, they did not transfer that habituation between stress types; for example, the clams that saw a shadow again and again would still react strongly to a different stress like tapping its shell. This suggests the animal can distinguish between different threats along a spectrum of seriousness, with touching of tissue (similar to a fish pecking at its flesh) being the most serious threat with the most dramatic response.

Another study determined that larger giant clams stayed closed longer than smaller ones in response to the same threat. They proposed this was related to the greater risk large clams face as they have more tissue area vulnerable to attack. While the clams might not have made a “conscious” decision in the way we do as thinking creatures, they were able to place their individual risk in context and vary their response. This ability to tailor a response to different risk levels is a sign of surprisingly complex neurology at work.

Inside the Scallop
Close up of the eyes of a scallop. Each is a tiny crystalline parabolic mirror (photo by Matthew Krummins on Wikipedia)

Scallops show some of the most complex bivalve behaviors. This relates back to their unique adaptations, including simple eyes that can resolve shapes and the ability to swim away from danger. Scallops have been found to discern between predator types by sight alone, to the extent that they did not initially recognize an invasive new predatory seastar as a threat. When swimming, they are capable of using this vision to navigate to places where they can hide, such as seagrass beds. It would be very interesting to compare the behavior of scallops in marine protected areas to those that can be freely harvested. Do they vary their behavior in response?

I hope I’ve made clear that while clams are not exactly intellectual powerhouses, their behavior is much more complicated than simply sucking up water and opening or closing their shells. Like us, they inhabit a complex environment that requires a multitude of responses. Their nervous systems have evolved to allow them to survive and adopt nuanced behaviors which they can vary on the fly, and which us “higher” animals are only just beginning to comprehend.

The Snails that Farm

File:Marsh periwinkle 001.jpg
Littoraria grazing on Spartina marsh grass. (source)

Us humans really like to talk up our skills at farming. And while it’s true that we have domesticated animals and plants to a degree not seen in other life forms, the act of nurturing and harvesting food is actually not really that special, and is broadly observed throughout the animal kingdom. Perhaps the most iconic invertebrate farmers are insects. Leaf-cutter ants, termites, and some beetles have been observed to actively cultivate fungus by gathering plant material to feed it, growing the fungus, protecting the fungus from competition, and then harvesting the fungus to feed themselves and their young. Ants are also known to keep livestock in the form of aphids, which they lovingly protect and cultivate for the sweet nectar they excrete. Such practices are called “high-level food production” because, like human farmers with their seeds and fertilizer, insects have evolved a highly complex symbiosis with their fungus. The fungus has shaped their behavior as much as the ants cultivate the fungus.
The marsh periwinkle Littoraria irrorata (source)

Less well understood is the “low-level food production” that may occur more broadly throughout the tree of life. There is less direct evidence of such behavior because it is more indirect and less specialized than high-level food production, but it may be equally advantageous for the cultivator and the cultivated. One study published in 2003 uncovered a simple but powerful relationship between marsh periwinkles of the genus Littoraria and fungus which they cultivate and harvest.
Close-up of a snail’s radula (source)

Marsh periwinkles are small and not particularly charismatic creatures. Like many snails, they are grazers with a shell, a fleshy foot and a rough, abrasive organ called a radula which they use like sandpaper to graze on pretty much whatever they can get into. Snails are not known as picky eaters. But researcher Brian Silliman of Brown University and Steven Newell of University of Georgia noticed that these innocuous snails regularly undertake the risky, low reward activity of grazing above the water on the blades of swamp grass, stripping off the surfaces of the blades of grass. The researchers were confused why the snails would expose themselves to predation and the harmful open air for such a low-nutrition food.


A typical snail farm, complete with liberally applied fertilizer. Yum.

They discovered that the snails were investing in the future. By stripping away the protective surface of the swamp grass blades and liberally fertilizing the surface of the grass with their droppings, the periwinkles are ensuring that the swamp grass will be infected with an active and very prolific fungal infection. The fungus, unlike the plant it lives on, is of high nutritional value. The researchers demonstrated the active partnership between the snails and fungus by conducting caged experiments where they showed that snails which grazed on grass but not the resulting fungus did not grow as large as snails which were allowed to return and chow down on the fungus. The fungus loves this deal as well. They grow much more vigorously on grass that is “radulated” (rubbed with the snail’s sandpapery radula) than uninjured grass. The fungus grows even faster if the snails are allowed to deposit their poop next to the wounds. The researchers found that this same relationship applies at 16 salt marshes along 2,000 km of the Eastern Seaboard.

The periwinkles don’t really know what they’re doing. They aren’t actively planting fungus and watching proudly like a human farmer as their crop matures. But over millions of years, the snails have been hard-wired to practice this behavior because it works. Snails that abrade a leaf of swamp grass, poop on the wound and come back later to eat the yummy fungus do a lot better than snails which just stick to the safe way of life below the surface of the water. The fungus loves this relationship too. The only loser is the swamp grass, which the researchers unsurprisingly found grows much more slowly when infected with fungus. But marsh grass is the largest source of biomass in swamp environments, and the snails that partner with fungus are able to more efficiently use this plentiful but low-nutrient food source, to the extent that it is now the dominant way of eating for swamp periwinkles on the East Coast of the US, and probably in a lot more places too. The researchers noted that there are likely far more examples of low-level food production that we simply haven’t noticed.

Since this work was published, other teams have discovered that some damselfish like to farm algae, fiddler crabs encourage the growth of mangrove trees, and even fungus get in on the action of farming bacteria. We love to talk up our “sophisticated” high-level food production techniques, but such relationships probably got started at a similarly low level. Our activities as hunter-gatherers encouraged the growth of certain organisms, we stumbled upon them, ate them, kept doing what we were doing and eventually our behavior developed into something more complex. Next time you see a snail munching its way up a blade of grass, consider to yourself whether it knows exactly what it’s doing. Come back later to see the fruits of its labor.

When a clam has a stowaway

My mussel contained a tiny half-eaten crab! - Imgur
Source: jeredjeya on Reddit

Bivalves put a lot of energy into their shells. These hardened, hinged sheaths of carbonate are an effective defense against many predators looking to get at the squishy clam’s body encased inside. Parasitic pea crabs have evolved to free-ride on the bivalves’ hard work.


(video courtesy Dana Shultz)

Pea crabs are small (pea-sized), very specialized parasites which live in the mantle cavity of many bivalve groups including oysters, mussels, clams and more. The mantle is the wall encasing the soft body of the bivalve, and the cavity is the space between this soft gooey tissue and the shell itself.

For a pea crab, there is no better place to be than this tiny, claustrophobic space. In fact, they can’t live anywhere else, though some species have been found in other unusual places such as inside the anuses and respiratory tracts of sea cucumbers (link SFW, fortunately, unless you’re a sea cucumber). In a bivalve host, the crab is protected from predation by the shell, and the bivalve provides a constant buffet of food as it sucks in suspended particles with its gills. The crab steals some of this food from itself before the bivalve can digest it.

As you might imagine, having a crab living in you taking your food and pinching at your gills is not an ideal arrangement for the bivalve. Pea crabs damage their hosts’ gills with their constant picking, and bivalves infected with crabs suffer slower growth than uninfected individuals, particularly for those unlucky enough to play host to the larger female pea crabs. At a certain point, the males will sneak out of their hosts and find a bivalve with a female crab inside. At this point, they mate inside the host’s shell, adding great insult to injury. The female releases her larvae, which swim out to infect new hapless bivalves and start the cycle over again.

Aww, she’s expecting! The papers refer to this as being “with berry” which I find amusing for some reason. (photo from Dana Shultz)


You might think that commercial oysters with crab parasites would be thrown out, but to the contrary, finding a pea crab or its close relative the oyster crab with your meal is a cause for celebration in some areas, such as the Cheasapeake Bay. The crabs are eaten whole and often raw, and are said to have a texture akin to shrimp, with notes of sweetness and umami. Personally I prefer surprises in my Kinder eggs rather than in my shellfish, but to each their own.

A pea crab serves as a nice side dish for this lunching sea otter. Source: Brocken Inaglory on Wikipedia


The Many Homes of Hermit Crabs

My boy Harry, a purple pincher (Coenobita clypeatus) inhabiting a tapestry turban snail (Turbo petholatus) shell. These seem to be his favorite kind, even though they do not come from his native Caribbean.

Hermit crabs (superfamily Paguroidea) are most famous for using snail shells as their home, having evolved a soft, spiral abdomen to be able to use them for protection. But they are more flexible about their choice of abode than you might expect.

This crab was likely preserved buried alive in sediment. Note how it uses its claw as a protective trap door sealing the opening of the ammonite shell. Source: Jagt et al, 2006

Different groups of shelled organisms have risen and fallen in abundance through geological time. During the time of the dinosaurs, ammonites (relatives of modern squid and octopus) were among the most common marine organisms, and hermit crabs were there to recycle their shells when they died.

Each tiny pore (zooid) in this bryozoan contained a tiny tentacled organism. Together they grew in a shape that made for a nice hermit crab house (image 5 shows a cross section where the crab’s abdomen would fit). Source: Taylor and Schindler 2004

Mollusks aren’t the only contractors for hermit crabs. Some hermits utilize the skeletons of colonial organisms like bryozoans as a home. Bryozoans are filter-feeding colonial animals made up of thousands of tiny tentacled organisms living in the pores of a shared skeleton. The extinct bryozoan Hippoporidra lived in symbiotic partnership with hermit crabs, growing around a gastropod shell to attract a hermit crab partner. This was an example of mutualism: by providing a home for a crab, the bryozoan would be transported to new environments with plentiful food particles to eat, and also would be protected from their arch-enemy, nudibranchs (sea slugs). Some modern day hermits, such as Manucomplanus varians of the Gulf of California, have evolved very similar partnerships with live staghorn corals.

Manucomplanus varians at Monterey Bay Aquarium

Not all hermit crabs live in hard houses. Some deep sea forms partner with anemones, with the stinging tentacles serving as an effective defense.

Source: Okeanos Explorer

The recently discovered green-eyed hermit crab, which also lives in deep water, lives in a glued-together mass of sand created by tiny anemones, which continue to grow the structure to fit the crab as it increases in size.

The green-eyed hermit crab was found over 200 m deep off of South Africa. Source: Lannes Landschoff via Eurekalert 

Unfortunately, hermits adapted for gastropod shells are unable to find adequate homes in some areas, due to overharvesting of shells for the tourist trade as well as an excess of plastic trash. These crabs make do with whatever items that they can find. Plastic is not an ideal home material for hermits. Bottlecaps and narrow tubes do not allow the crab to fully retract for protection and leach chemicals which may harm the crab. The crabs also nibble on their shells as a source of calcium, which is obviously not possible with plastic.

Coenobita purpureus, a land hermit crab on Okinawa. Source: Shawn Miller

But hermits continue to impress me with their flexibility and ingenuity in their search for homes. For a hermit crab, home is where the abdomen is.