"So, naturalists observe, a flea has smaller fleas that on him prey; and these have smaller still to bite ’em; and so proceed ad infinitum."
- Jonathan Swift

February 16, 2012

Acanthocephalus dirus

The word parasite has a lot of connotations associated with it, and "maternal" is certainly not one of them. To most people, the term "freeloader" comes to mind (hopefully, this blog will show you that parasitism is actually a very challenging way of life). They also have a reputation as being pretty lousy parents. In most textbooks, parasites are usually considered as "r-strategists" - which produce many, many offspring and don't take good care of them (as opposed to a K-strategist which produces fewer offspring, but invest a lot into parental care - like an elephant). But not all parasites are bad parents, and today, I am going to tell you about a study on a maternal parasite which sacrifices everything (literally) for her offspring.

Acanthocephalus dirus has a reproductive strategy that is unusual for its group - the acanthocephalans or the thorny-headed worms (Acantho = "thorns", Cephala = "head"). In fact it is unusual compared to most intestinal parasites. Unlike some tapeworms, which profligately cast off segments (each containing hundreds of eggs) into the wilderness with abandonment, A. dirus has rather different approach. The impetus that spurred on this piece of research were two separate observations: (1) fish that are infected with A. dirus do not have any worm eggs in their feces (unlike most animals infected with intestinal parasites) and (2) perfectly healthy and intact female worms were often expelled from the definitive host. What the researchers found was that instead of simply laying eggs that are expelled from the worm and from the host, a female A. dirus actually retains her eggs until she become completely bloated with them - at which point she exits gracefully from the host fish's digestive tract. Some readers might recall a nematode that has a similar reproductive strategy, and that both lineages have evolved such a reproductive strategy independently. So why has A. dirus evolved such an extreme strategy instead of just laying eggs normally like other thorny-head worms?

One reason could be that A. dirus infects creek chub - which, as its name indicates - lives in flowing creeks. The chub acquire the worm through eating infected isopods in the stream (the picture shows the light-coloured infected isopod on the right, and the darker uninfected individual on the left), which become infected when they ingest worm eggs resting on the creek bed. Acanthocephalan eggs tend to float - so if the eggs are simply expelled into the environment, they would get washed away downstream and deposited where the isopods do not occur. Whereas with A. dirus, the worm's own body can act like a weight belt which would carry the eggs down to the sediment layer, so by the time the worm herself decays, the eggs are already in the sediment where isopods can pick them up.

Furthermore, laboratory tests showed that isopods like to eat egg-filled female worms as much as their usual food - leaf litter - and the worm body itself actually enhances the infection success of the eggs. Researchers found that when exposed to fresh eggs alone, fewer than one in four isopods became infected, whereas when exposed to gravid females, over 80% became infected (natural infection comes somewhere in between those at about 60%). By making the ultimate maternal sacrifice, A. dirus gives her offspring the best possible start in life.

Image from figure in: Seidenberg (1973) Journal of Parasitology 59: 957-962

Kopp, D.A., Elke, D.A., Caddigan, S.C., Raj, A., Rodriguez, L., Young, M.L. and Sparkes, T.C. (2011) Dispersal in the acanthocephalan Acanthocephalus dirus. Journal of Parasitology 97: 101-105

February 9, 2012

Ascarophis sp.

When I saw the reports of giant amphipods being dragged up from the Kermadec Trench off the coast of New Zealand, my immediate thought was "I wonder what parasites it has?" This promoted me to do a write-up of a paper I've read recently, which is about a parasite that infects amphipods - admittedly those that are more modestly sized. Today, we are featuring a study on Ascarophis, a nematode worm that infects an intertidal amphipod (Gammarus deubeni) in Passamaquoddy Bay, New Brunswick, Canada. Compared with related species this worm has evolved to live the simple life(-cycle), and avoids the complications that come with having a complex life-cycle.

Previously on this blog, we have featured parasites that have evolved to take short-cuts with their complicated life-cycles. When a particular host is absent, such parasites may opt to ditch that host from their life-cycle, and switch up their developmental schedule. This is the case with the fluke Coitocaecum parvum. However, while C. parvum can switch between different life-cycles depending on circumstances, Ascarophis has completely abandoned that altogether, and has evolved to make things simpler by completing its entire life-cycle within its amphipod host. Usually, parasites with complex life-cycles use different hosts for different functions - i.e., one host might merely serve as a transport and/or resources for temporary development, whereas another acts as a mating ground and/or habitat in which it reaches maturity. So how can Ascarophis get so much functionality out of a tiny little crustacean?

Nematodes normally go through 4 larval stages (L1-L4) before becoming a sexual mature "fifth stage" worm (L5). The end of each larval stage is accompanied by a molt (rather like insects). In related nematodes that have retained their complex life-cycle, the L3 worms (which are ready to infect the next host) live encapsulated in the first host, while the L4-L5 live in the digestive tract of the final host. What the researchers found with the Ascarophis they collected from New Brunswick is that L1 and L2 worms were found in the muscle tissue, and upon reaching L3 the worms begin to migrate into the body cavity where they complete their development into adulthood and start producing eggs. Now compare this with Ascarophis from the White and Baltic Seas, which also infect amphipods, but uses a species of sculpin as their final host. Those fish acquire their infection by eating amphipods infected with L3 stage nematode, and the worms develop into adults in the fish's gut.

In effect, the Ascarophis from New Brunswick gets the most out of its little crustacean host by using different parts of the amphipod's body as surrogates for different hosts - instead of being transmitted to a different host, it simply moves to occupy a different part whose function is close enough to its needs for it to complete its development. Unlike the C. parvum, it appears that Ascarophis has abandoned the fish host altogether, and has committed itself to using the amphipod as the sole host for its entire life-cycle. Even though the Ascarophis found in the White and Baltic Seas have retained their complex life-cycle, researchers of this study suggested that they are the same species as the worms they looked at, but the New Brunswick variant has simply adapted to local condition and evolved a different life-cycle. However, it must be noted that the researchers have come to this conclusion based on the worm's morphology and as we have seen before, appearance can be deceptive with nematodes.

Through all that, this plucky little New Brunswick parasite faces one last problem - getting its eggs out of its crustacean host. For worms that live in inside a fish's gut, passing eggs out into the environment is a pretty straightforward affair - the eggs simply get washed out with the poop. But there is no exit in the body cavity of an amphipod, so how is a worm supposed to cast its eggs out into the environment? Well, this thrifty nematode simply waits for the host to die, and as the body disintegrates, the eggs are released as well. Of course, it helps that these amphipods have a tendency to cannibalise the rotting bodies of their fallen comrades - this presents the perfect opportunity for the parasite to infect a new batch of hosts - yet another reason to not gnaw on any random corpses you may come across.

Image modified from figure in the paper

Appy, R.G. and Butterworth, E.W. (2011) Development of Ascarophis sp. (Nematoda: Cystidicolidae) to maturity in Gammarus deubeni (Amphipoda). Journal of Parasitology 97: 1035-1048.