Revenge of the Clams

Lampsilis showing off its convincing fish-like lure. Photo: Chris Barnhart, Missouri State.

Clams are traditionally the victims of the aquatic realm. With some exceptions, clams are generally not predatory in nature, preferring to passively filter feed. When they are attacked, their defenses center around their protective shell, or swimming away, or just living in a place that is difficult for predators to reach. They are picked at by crabs, crushed in the jaws of fish, and pried apart by sea stars. But some clams are sick of being the victims. They have big dreams and places to be. For these clams, the rest of the tree of life is a ticket to bigger and better things. These clams have evolved to live inside of other living things.

Pocketbook mussels, for example, have a unique problem. They like to live inland along streams but their microscopic larvae would not be able to swim against the current to get upstream. The mussels have adapted a clever and evil strategy to solve this problem: they hitch a ride in the gills of fish. The mother mussel develops a lure that resembles a small fish, complete with a little fake eyespot, and invitingly wiggles it to attract the attention of a passing fish. When the foolish fish falls for the trick and bites the mussel’s lure, it explodes into a cloud of larvae which then flap up to attach to the gill tissue of the fish like little binder clips. They then encyst themselves in that tissue and feed on the fish’s blood, all the while hopefully hitching a ride further upstream, where they release and settle down to a more traditional clammy life of filter-feeding stuck in the sediment.

Very tiny Mytilus edulis living in the gills of a crab (Poulter et al, 2017)
The tiny 2.5 mm long Mimichlamys varia, living on the leg of a crab (Albano and Favero, 2011)



Clams live in the gills of all sorts of organisms. Because they broadcast spawn, any passing animal may breathe in clam larvae which find the gills a perfectly hospitable place to settle. Sure, it’s a bit cramped, but it’s safe, well oxygenated by definition and there is plenty of food available. They also may just settle on the bodies of other organisms. Most of these gill-dwelling clams are commensal: that means that their impact on the host organism is fairly neutral. They may cause some localized necrosis in the spot they’re living, but they’re mostly sucking up food particles which the host doesn’t really care about. In addition, in crabs and other arthropods, these clams will get shed off periodically when the crab molts away its exoskeleton, so they don’t build up too heavily.

Top: Kurtiella attached among the eggs of the mole crab. Bottom: aberrant Kurtiella living within the tissue of the crab (Bhaduri et al, 2018)

While being a parasite is often denigrated as taking the easy way out, it is actually quite challenging to pursue this unusual lifestyle. Parasitism has evolved a couple hundred times in 15 different phyla, but it is rare to find some organism midway in the process of becoming a true parasite. One team of researchers just published their observations of a commensal clam, Kurtiella pedroana, which may be flirting with true parasitism. These tiny clams normally live in the gill chambers of sand crabs on the Pacific coasts of the Americas. They attach their anchoring byssal threads to the insides of the chambers and live a comfortable life until the crabs molt, when they are shed away. The crabs mostly are unaffected by their presence, but the researchers noticed that some of the clams had actually burrowed into the gill tissue itself. This is an interesting development, because the clams would not be able to filter feed in such a location, so they must have been feeding on the crab’s hemocoel (internal blood). These unusual parasitic individuals are currently a “dead end” as they haven’t figured out how to get back out to reproduce, but if they ever do, they could potentially pass on this trait and become a new type of parasitic clam species. The researchers have potentially observed a rare example of an animal turning to the dark parasitic side of life, with some living in a neutral commensal way and other innovative individuals seeking a bit more out of their non-consensual relationship with their host crabs.  Considering the irritation that other bivalves suffer at the claws of pesky parasitic crabs, this seems a particularly sweet revenge.





Hard shells aren’t actually that hard to make (yet)

One of the Antarctic bivalve species featured in this study. Source

Like all organisms, bivalves have a limited budget governing all aspects of their metabolism. If they put more energy into feeding (filtering the water), they can bring in a bit more food and therefore fuel more growth, but sucking in water takes energy as well, particularly if there isn’t enough food to be filtered out. Bivalves also periodically have to grow gonadal material and eggs for reproduction, expand their body tissue (somatic growth) and of course, grow their shells (made of of a mineral called carbonate). All of these expenditures are items in a budget determined by the amount of energy the bivalve can bring in, as well as how efficiently they can digest and metabolize that energy.

If a bivalve is placed under stress, their scope for growth (the max amount of size increase per unit time) will be decreased. Because they’re cold-blooded, bivalves are limited by the temperature of their environment. If temperatures are low, they simply can’t sustain the chemical reactions required for life at the same rate that endotherms like us can. They also may have to shut their shells and stop feeding if they’re exposed by the tide, or are tossed around by a violent storm, or attacked by predators or toxins from the algae that they feed on.

When their budget is lower, they have to make painful cuts, much like a company lays off employees if their revenues are lower. The question is which biological processes get cut, and when? My first chapter (submitted and in review) has settled temperature being the primary control on seasonal shell growth. Bivalves at high latitudes undergo annual winter shutdowns in growth, which create the growth bands I use to figure out their age, growth rate, etc. We’d be a lot closer to accurately predicting when bivalves suffer from “growth shutdowns” if we had hard numbers on how much energy they actually invest in their shells. A new study from a team led by Sue-Ann Watson of James Cook University attempts to do just that.

Diagram relating the growth bands of Antarctic soft-shelled clam with a chart showing the widths of those bands. Source

Collecting a database of widths for the annual growth rings of bivalve and gastropod (snail) species from many latitudes, Watson and her team were able to get a global view of how fast different molluscs grow from the equator to the poles. Because the unit cost of creating carbonate is determined by well-understood chemistry, they were able to create an equation which would determine the exact number of Joules of energy used for every bivalve to grow their shells.

They still needed a total energy budget for each species, in order to the percent of the energy budget that each bivalve was investing in their shells. They drew on a previous paper which had calculated the standard metabolic rates for each species by carefully measuring their oxygen consumption. We could do the same for you if you sat in a sealed box for an extended period of time while we measured the exact amount of oxygen going in and CO2 going out. Dividing the amount of energy needed to grow the shell by the total amount of energy used in the organism’s metabolism would give us a percent of total energy that the bivalve dedicates to adding growth layers to its shell.

That number is…not very large. None of the bivalves or gastropods they looked at put more than 10% of their energy into shell growth, and bivalves were the lowest, with less than 4% of their energy going into their shell. Low-latitude (more equatorward) bivalves have the easiest time, putting less than 1% of their energy into growth but getting way more payoff for that small expenditure. High-latitude polar bivalves have to work harder, because the lower temperatures they experience mean the reactions needed to create their shells are more expensive. In addition, most of that energy is going into the protein-based “scaffolding” that is used to make the shell, rather than the crystals of carbonate themselves. Organisms right now don’t have to put a whole lot of effort into making their protective shells, which could explain why so many organisms use shells for protection. That is good, because if shells were  already breaking the bank when it came to the bivalves’ growth budget, they wouldn’t have a lot of room to invest more energy in the face of climate change. Unfortunately, as the authors note, these budgets may need to change in the face of climate change, particularly for bivalves at the poles. As the oceans grow more acidic due to human CO2 emissions, growing their shells will start to take up more of their energy, which is currently not a major part of their budget.

A cold-water ecosystem dominated by Antarctic scallops. Source

Right now, the cold waters of the poles are refuges for organisms that don’t deal well with shell crushing predators. As polar regions warm, such predators will begin to colonize these unfamiliar waters. Polar bivalves may encounter the double whammy of needing to spend more energy to make the same amount of shell, but also find that it is no longer enough to protect them from predators that easily crack open their protective coverings.

I found this study to be an elegant and thoughtful attempt to fill in a gap in our current understanding of how organisms grow and how energy budgets are influenced by environmental variables like temperature. I instantly downloaded the paper because it answered a question that has long been on my mind. Maybe can sneak its way into my manuscript during the review process!