Feeding or sex (or both)?

About three weeks ago I collected some mussels from the intertidal, to use both in the lab and in the classroom. A mussel can itself be an entire habitat for many other organisms. Many of the mussels I brought into the lab this last time were heavily encrusted with barnacles and anemones. I wanted to look more closely at one of the anemones so I took the mussel to the microscope. And, as often happens when I look at stuff under the microscope, I got totally distracted by things other than what I intended to.

But for the record, this is the anemone that started the whole chain of events:

A small aggregating anemone (Anthopleura elegantissima). 5 December 2016 © Allison J. Gong
A small aggregating anemone (Anthopleura elegantissima).
5 December 2016
© Allison J. Gong

Below the anemone there's a thick mat of small acorn barnacles (Balanus glandula) and a couple of leaf barnacles (Pollicipes polymerus). They were all alive when I brought the mussel into the lab, and over the weeks a few of them have died. But many of them are still kicking, both figuratively and literally.

Barnacles are most strange animals. Believe it or not, they are crustacean arthropods, somewhat closely related to crabs and lobsters. They live encased within a shelter of calcareous plates, which they can close seal up against predators and desiccation. I've never figured out why they are called "acorn barnacles," as they don't look anything like acorns to me, but in Balanus and such the base of the shelter is glued directly to a rock or some other hard surface. Leaf barnacles are shaped very differently, and have a fleshy stalk between the shelter that houses the main body of the animal and the rock surface.

Small acorn barnacles (Balanus glandula). 5 December 2016 © Allison J. Gong
Small acorn barnacles (Balanus glandula).
5 December 2016
© Allison J. Gong
Close-up view of a leaf barnacle (Pollicipes polymerus). 5 December 2016 © Allison J. Gong
Close-up view of a leaf barnacle (Pollicipes polymerus).
5 December 2016
© Allison J. Gong

To picture what's going on with a barnacle, imagine a shrimp lying on its back, then curl it up and stick the whole thing inside some calcareous plates. The thoracic appendages would be facing up. In barnacles the thoracic appendages are modified to be clawlike feeding structures called cirri. Barnacles are filter-feeders, collecting particles from the water by maneuvering the cirri in a sort of grasping fashion. So in a nutshell, or more precisely a test, a barnacle lies on its back and kicks its legs out to catch food.

Here's what B. glandula looks like when feeding. Note the clearly jointed cirri, with fine hairs that help catch particles. The cirri can be controlled independently, as you can see when they flick towards the center, and the entire apparatus can be rotated quite a bit.

Same deal with Pollicipes.

So that's the feeding part. A little strange, but not as interesting as the barnacles' sex lives. Let's start with some background about sexual function. And get your mind out of the gutter; this is real science stuff! Most of the animals that you're familiar with are described as dioecious (Gk: "two houses"). This means that female and male sexual functions are segregated; in other words, there are male bodies and female bodies. Other animals are described as monoecious (Gk: "one house"), so that a single body has both female and male sexual function. Monoecious animals could also be described as hermaphroditic. Some monoecious animals have male and female function in a single body at the same time; we call these simultaneous hermaphrodites. If a body first functions as one sex and then either acquires or switches to the other sex, we say the animal is a sequential hermaphrodite. Many fishes, including the California sheephead and the anemone fishes of coral reefs, are sequential hermaphrodites. Make sense?

Barnacles are simultaneous hermaphrodites. If you dissect an adult barnacle you will find mature ovaries and testes. This means that every barnacle can be both a mother and a father. The logical assumption is that monoecious animals should just fertilize their eggs with their own sperm. . . however, this generally isn't the case. The whole point of sexual reproduction is to combine the genomes of two individuals, and self-fertilization obviously doesn't accomplish this. So even though there are many hermaphroditic animals, very few of them are self-fertile.

One other weird thing about barnacles, and crustaceans in general, is their sperm. Arthropods have non-flagellated sperm, which means they don't swim (although some of them have amoeboid sperm that can ooze around a little bit). Many marine animals reproduce by broadcast spawning; that is, by throwing their gametes into the water, where fertilization takes place. Fertilization is facilitated by the sperms' ability to swim towards conspecific eggs.

Barnacles, with their non-swimming sperm, generally cannot rely on broadcast spawning to get sperm to egg. They must copulate. How do you suppose they do this? The same way that other animals (e.g., Homo sapiens) copulate, by using a penis or some other structure to transfer sperm from one individual to the body of another. In barnacles the penis's technical name is intromittent organ. The penis is inserted into the test of a neighboring barnacle and sperm is delivered. The receiving barnacle uses the sperm to fertilize its eggs. Unlike the cirri, the penis is unjointed and flexible, the better to seek out and slip into potential mates. You can see the intromittent organ unrolled and poking around.

Now think about the ramifications of these constraints. Barnacles live their entire post-larval lives permanently cemented to a rock. They also have non-motile sperm so sperm transfer can occur only by copulation. If the key to reproductive success is to mate with as many other individuals as possible, what do you suppose natural selection has done? That's right: barnacle anatomists, including the great Charles Darwin himself, have noticed that barnacles have incredibly long penises. In fact, compared to overall body size, barnacles have the longest penises in the animal kingdom, up to 15 times the length of the body! That's what you call bragging rights. Not all barnacle species are so amply endowed, however. The same leaf barnacle that I observed today (P. polymerus) has recently been reported to be a spermcaster; their penises are shorter than body length, and they release sperm that are captured by their downstream neighbors.

Wonders never cease.

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