Finally! At long last I have evidence that my juvenile urchins have mouths and are feeding. A week ago I put a batch of seven teensy urchins onto a scuzzy glass slide and have been watching them daily ever since. And yesterday, just as I was beginning to worry that they’d never be able to eat, I saw that some of them had eaten little tracks through the scuzz on the slide.
The little urchin still has a test diameter of about 0.5 mm, so it hasn’t really started growing yet. However, see the squiggly dark paths? Those are areas of the slide that have been eaten clean. The scuzz is algal in origin, giving the slide an overall brownish-green color, so the scuzz-free parts of the slide are clear–or dark, actually, given that I took this photograph against a black background–having been munched clean by the urchin’s teeth. And the other bit of evidence that I saw? Poop! Yes, there were fecal pellets on the slide, which proves that the little urchin has a complete functional gut.
And those small round golden objects you see on the slide? Those are big centric diatoms of the genus Coscinodiscus. They are the only local diatoms that I know of that are big enough to be seen with the naked eye.
Lastly, because I just can’t seem to stop myself, here’s a video of the little urchin:
I love the sculpturing of the spines. And do you see that three-pronged structure at about 9:00 on the urchin? That’s a pedicellaria. On adults of the genus Strongylocentrotus there are four types of jawed pedicellariae, three of which, in my experience, are easy to distinguish on a living specimen. But in this young an animal I can’t yet tell how many types of pedicellariae it has. I suppose that the formation of pedicellariae might be the next event for me to follow as these urchins continue to grow and develop.
Everybody knows that climate change is a hot–pun intended!–topic in both science and politics these days. Here along the northern California coast it seems that sea surface temperature (SST) has been elevated for at least a year now. I remember a time, not too many years ago, when I would put my hands into my seawater table and they’d go numb after several minutes. This told me that the water temperature was in the 11-12 ºC range. But that hasn’t happened for a while, and recently I’d put my hands in the water and it didn’t even really feel cold. My trusty not-fancy thermometer has been telling me that the temps have been hovering at around 14ºC.
The other day it occurred to me that I have a 20-year record of water temperatures from my seawater table, which is a pretty fair proxy for SST in the area. The numbers may not jive exactly with SST data produced by oceanographic instruments, but the trends should be very similar. If you click on the figure you’ll be able to see a larger version of it.
There are a couple of notable trends in these data. I was pleased to see a strong signal for the 1997-1998 El Niño event, visible as a prolonged period of elevated temperatures in the fall and winter. This was followed by a La Niña in 1998-1999, when temperatures were lower than average for a few months. Aside from those events, SST fluctuates between about 16º in the summer-fall and 11-12º in the winter-spring.
One more thing. Take a look at the far right end of the graph. Notice what appears to be a cooling trend so far in the spring of 2015?
Here are the data from March and the first three days of April:
So there’s definitely a cooling trend in the past few days. The interesting question is: Why is this happening now, when it hasn’t happened for about two years?
The answer, in a nutshell, is the wind. For the past week or so, we’ve had screaming afternoon winds at the marine lab, coming from the northwest. Northwest winds blowing down the coast drive the process of coastal upwelling, which results in cold water rising to the surface; it usually takes 3-4 days of sustained winds to start upwelling. This upwelled water, in addition to being cold, also contains a lot of nutrients, which are used as fertilizers by the primary producers of the marine ecosystem, the phytoplankton. Most of the phytoplankton are photosynthetic unicellular algae (NOT plants) that harvest the energy from sunlight and use it to fix carbon dioxide into organic molecules. The fixed carbon in turn feeds grazers such as copepods, which are then eaten by small predators, which are eaten by larger predators and so on up the food chain.
What this all means is that we may, for the first spring in two years, be getting some productive upwelling. I don’t think I’m the only marine biologist in the area who is looking forward to seeing whether this apparent upwelling continues. If it does, then we should see the biota respond accordingly. Mind you, a four-day streak does not indicate a long-term return to typical spring upwelling conditions, and it may be merely a blip in the warmer conditions that are the new normal for us, but it is a stronger signal than we’ve seen in a few years. In any case, I will be keeping an eye on both the water temperature and the critters living in it.
Anyone who went to graduate school in the sciences remembers what oral exams are like. I remember not having any fun at all in mine, and by the time I was dismissed I wasn’t sure what my own name was. Fortunately, that is all ancient history and now I get to spend my time performing a different kind of oral examination on other creatures.
My oldest urchins are now 17 days post-metamorphosis and I’ve been watching them to see when their mouths break through. It seems to me that 17 days is a long time, but the time is near. Besides, the animal is always right. In the urchin that I examined closely the five teeth of Aristotle’s lantern are very close to breaking through the thin membrane covering the mouth opening. The teeth are also much more active than they were a week earlier, as you can see in this short video clip:
I also checked out another tiny urchin and noticed that this individual has startlingly red buccal tube feet:
Sea urchins have five pairs of large tube feet on the oral surface, surrounding the mouth. As with all tube feet, the buccal tube feet are part of the animal’s water vascular system and are situated in the ambulacral region of the test; they are used to manipulate and grab food. In adults of this species, the buccal tube feet are much larger and more robust than the other tube feet. In this little guy the tube feet are noticeably red, but I can’t yet tell if they’re bigger than the others.
And just for kicks I took another video:
Yesterday I transferred seven urchins onto a glass slide that I’ve had basking in the sun in an outdoor tank to develop a thin film of algae. As the urchins’ mouths become functional they should be able to start munching on the scuzz on the slide. So far they seem happy to be crawling around on the slide but this morning I didn’t see any signs that they’d actually eaten anything.
My oldest baby urchins have been actual sea urchins for eight days now. Their total age, counting from the time they were zygotes, is 58 days. When an animal undergoes a life history event as drastic as this metamorphosis, it can be tricky deciding how to determine its age. Do you count from when egg and sperm formed the new zygote, or from when the juvenile (and eventually adult) body form was achieved? For the sake of this discussion I’m going to count from the date of fertilization, simply because I know exactly when that date was and it’s the same for all of these larvae, larveniles, and juveniles. This just makes sense to me.
So, at the grand old age of 58 days, which is five days post-metamorphosis for the oldest individuals, the baby urchins have grown a lot more tube feet, spines, and pedicellariae. However, they haven’t gotten any bigger. This is because they aren’t eating yet. I’ll explain why in a bit. The individual in the picture below measures about 490 µm in test diameter–that’s the opaque part in the center of the animal. The spines make the apparent size much larger.
In this short video clip you can see how many more tube feet this animal has, compared to the original five it started with. The movements are now much more coordinated, too, and these animals can walk with what appears to be purposeful direction. You can also see the texturing of the spines and the little pincher-like pedicellariae.
To see the surface details of the animal when it’s this opaque, I needed to use a different kind of lighting. Instead of using the transmitted light that shines through the object on the stage of the microscope, giving a brightfield view, I used my fiber optic light to create a darkfield effect that shows the surface details of the animal. Then I shot another video clip with this epi-illumination and focused up and down on the oral surface to see what was going on there. Fortunately the baby urchin isn’t yet able to right itself very quickly, and it stayed oral-side-up for as long as I needed to take the photos and video.
What this video clip of the urchin’s oral surface shows very clearly is that the animal doesn’t have a mouth yet. The pinkish star-shaped structure in the center is actually the negative space between the five triangle-shaped white teeth which all point to the middle. Soon, I expect in the next handful of days or so, that thin membrane covering the mouth will rupture, and the teeth will be exposed for the first time to the outside environment. At that point the urchin will begin feeding.
You may well be wondering, How the heck are they living if they haven’t eaten in over a week? They’re babies, after all, and don’t babies have to eat all the time? Well, yes, they are babies. But before they were baby urchins they were larvae, and as larvae were kept well fed by yours truly for their entire larval life. Part of becoming competent as a larva is sequestering enough energy stores to power the process of metamorphosis and keep the juvenile going until it has a mouth and can feed itself. Remember, this new animal has to do everything–locomote, eat, avoid predators–with body parts that it didn’t have when it was a larva. Building whole new body parts and learning how to use them takes time. So these newly metamorphosed juveniles have about 10-12 days to fast until their mouths break through and they can begin eating. Any individual that didn’t store enough energy to make it through the fast, will die.
I’ll check on them again tomorrow (day 59) and see if it’s time to transfer the juveniles to their food source, which will be algal scuzz that I’ve been cultivating on glass slides for a few weeks. They’ll grow quickly once they’re eating. I hope I have enough scuzz to keep up with them!
Imagine spending your entire life up in the water column as a creature of the plankton. You use cilia to swim but are more or less blown about by the currents, never (hopefully!) encountering a hard surface, and feeding on phytoplankton and other particulate matter suspended in the water. Then, several weeks into your life’s adventure, you fall out of the plankton, dismantle your body while simultaneously building a new one, and about a day later have to begin walking using anatomical structures that you didn’t have 24 hours earlier. Not only that, but the food that you’ve been eating your entire life is no longer available to you, for you no longer possess the apparatus that can capture it. And, finally, your body symmetry makes a wholescale change from bilateral to pentaradial–just think of what that means in terms of how your body is oriented and moves through three-dimensional space. That’s what metamorphosis is like for sea urchins and many other echinoderms.
The objects of my complete and utter obsession for the past month and a half have started metamorphosing from small larvae into tiny urchins. When I did my daily check yesterday I had two that had completed metamorphosis since the previous day. One of them still had a bit of puffiness on the aboral surface, which I think may be the very last remnants of the larval body. This little guy has only its first five tube feet, from the juvenile rudiment of the competent larva.
But just having feet doesn’t mean you automatically know how to walk with them, and it’s no easier for these guys than it is for humans. It’s probably more difficult, actually, because the urchins have to coordinate movement of five appendages simultaneously. They typically pick up one or two tube feet from the same side of the body and wave them around until one of them randomly sticks to something. Then they remain stretched out until the tube feet on the opposite side of the body let go. Well, you can watch for yourself; this is the same individual that is in the top photo above:
Being a bit farther along in the developmental process means having more spines, but not necessarily any more coordination. I watched the second urchin for several minutes, and while it repeatedly detached and re-attached tube feet, it didn’t actually go anywhere. Here’s a short clip:
It’s amazing how quickly they learn, though. When I go to the lab to look at them tomorrow, they’ll be running around as though they’ve been ambulatory their entire lives. Which, in a peculiar sense, depending on when you start counting, maybe they have.
After much teasing and titillation, my urchin larvae have finally gotten down to the serious business of metamorphosis. It seems that I had jumped the gun on proclaiming them competent about a week ago, or maybe they were indeed competent and just needed to wait for some exogenous cue to commit to leaving the plankton for good. In any case, I’ve spent much of the last five days or so watching and photographing the larvae to document the progress of metamorphosis as it occurs. While I was unable to follow any individual larva through the entire process of metamorphosis, I did manage to put together a series of photographs that document the sequence of events.
To recap: A competent larva is anatomically and physiologically prepared to undergo metamorphosis. This batch of larvae reached competence at the age of about 45 days. The larva below is very dense and opaque in the main body. It can still swim, but has become “sticky” and tends to sit on the bottom of the dish.
Sometimes the first tube feet emerge from the larva while it is still planktonic. Other times the larva falls to the benthos and lands on its (usually) left side, where the rudiment is located.
This larva is lying on its right side, so the tube feet are sticking straight up out of the plane of view. You can clearly see two of them, though.
Just for kicks, here’s the same larva, photographed with dark-field lighting. This kind of light illuminates the surface of the object being viewed, which is very helpful when the subject is opaque, making it possible to see four tube feet in this picture.
As the tube feet are emerging from the juvenile rudiment, the larval body contracts and gets denser. The arms shrink and the internal skeletal rods that supported them are discarded. At this stage the larval juvenile (larvenile? juvenal?) begins to crawl around on the bottom. The ciliated band that used to propel it through the water and create the feeding current may still be beating, but eventually will stop, as the larvenile will no longer need it. This is usually the time that I see the first spines waving around; it’s interesting to note that tube feet, which originate from the inside of the animal, come first, then are followed by spines. Then again, the spines are part of the animal’s endoskeleton, so maybe it’s not so noteworthy after all.
Anybody who has visited one of the sandy beaches in California has probably seen kids running around digging up mole crabs (Emerita analoga). These crabs live in the swash zone at around the depth where the waves would be breaking over your ankles, moving up and down with the tide. They are bizarre little creatures, burrowing backwards into the sand with just their eyestalks and first antennae reaching up into the water.
Although it’s called a mole crab, Emerita‘s external anatomy isn’t very similar to that of other crabs. For one thing, it doesn’t have claws. In fact, its legs are quite unlike the legs that you’d see in a typical crab. Check out Emerita‘s appendages:
External anatomy of Emerita analoga
The crab’s head faces to the left in this diagram. The real surprise that these little crabs hide is the nature of the second antennae. Usually the crab keeps these long, delicate antennae protected under its outer (third) pair of maxillipeds. This is why you don’t see them when you dig up a mole crab.
You do see them when the crabs are feeding. As a wave washes over the crab, it extends the second antennae and pivots them them around on ball-and-socket joints. The feathery antennae catch particles in the water, then are drawn underneath the maxillipeds so the food can be slurped off and eaten.
Here’s a top-down view of two Emerita feeding. The purple-grayish thing in the field of view is a sand dollar (Dendraster excentricus).
This side view gives a better angle of what’s going on:
I find these little crabs quite captivating. I love how they rise up when I put food into their tank. Watching them feed always makes me smile.
In the parlance of invertebrate zoologists, competence is the state of development when a larva has all of the structures and energy reserves it needs to undergo metamorphosis into the juvenile form. In the case of my sea urchins, this means that they have four complete pairs of arms, each with its own skeletal rod, and a fully formed juvenile rudiment, which contains the first five tube feet of the water vascular system. A continuous ciliated band runs up and down all eight arms and provides the water current used both for swimming and feeding. The larva will have been eating well and its gut will be full of food. It will have lost the transparency it had when it was younger and will appear to be more solid-looking in the central area.
The first batch of larvae that I began culturing this season are now 42 days old. Some of these are competent, or very nearly so. Last week I isolated about a dozen of these big guys into a small dish, making it easier for me to observe them closely every day. Today they looked decidedly opaque and dumpy, and although some of them were still swimming others were heavy and tended to rest on the bottom of the dish.
General orientation: This is a ventral view. The animal swims with its arms forward, which defines the anterior portion. Thus the bottom of the cup-shaped body is the posterior. This larva measures ~900 microns along the anterior-posterior axis. Plutei have bilateral symmetry that goes all to hell during metamorphosis, from which the urchin crawls away with typical echinoderm pentaradial symmetry. This wholescale change in body organization is one of the truly amazing things about metamorphosis in these animals. It boggles my mind every time I think about it.
You can see that this pluteus has eight arms. The oblong reddish structure in the middle is the stomach, which has taken on the color of the food the animal has been eating. The strange mixed-up looking structure adjacent to the stomach on the animal’s left side is the juvenile rudiment. Focusing up and down through the rudiment shows that it contains five tube feet. After metamorphosis, the juvenile urchin will use those first five tube feet to walk around as a benthic creature, having spent all of its life up to this point as a member of the plankton.
Today I captured about 20 seconds of a larva feeding. This individual is a day older than the one in the photo above and has more of that opacity that I associate with competence.
This is a dorsal view; if you imagine that you’re looking at the animal’s back, you see that the rudiment is indeed on its left side. The larva’s ciliated band is moving a lot of water, and the little specks that you can see flying around are food cells. There wasn’t enough water in this drop for the pluteus to do any actual swimming, but at this point it’s pretty heavy and would tend to sink to the bottom.
Some time in the next several days these guys are going to start metamorphosing. I will be examining them every day; keep your fingers crossed that I catch them in the act!
Thirty-one days ago, on 20 January 2015, I spawned purple sea urchins (Strongylocentrotus purpuratus) and generated six jars of larvae. I’ve been examining the larvae twice a week ever since. At first they were doing great, developing on schedule with no appreciable abnormalities or warning flags. Then, at about Day 24, the cultures began crashing for no apparent reason. At first I expected to see lots of malformed, shriveled, or underdeveloped larvae, but the thing that I don’t understand is that for the most part they look great. They’re eating, pooping, growing, and (apparently) doing everything that they should be doing.
This larva is PERFECT. It has all four pairs of arms now, and they are growing symmetrically. The stomach (the inverted-pear-shaped structure in the middle of the cup-shaped region) is pigmented with the red food it has been eating, and there are no skeletal rods protruding beyond the tips of the arms. This individual doesn’t give me any clues as to why the culture it came from took a nosedive this past week. The other larvae that I sampled from this jar today also look good. There aren’t many left in the jars from this spawning, but if they all look as promising as this one then I still have hope that some will be able to metamorphose successfully.
So what gives? I suspect that Day 24 has something to do with it. I’m working on a hypothesis and need to let it percolate inside my brain a bit more. When it’s ready I want to test it, although that will have to wait until next year, as we’ve reached the end of this year’s spawning season.
Yesterday I drove up the coast to Pigeon Point to do a little poking around. I had originally planned to search for little stars, survivors that had made it through the most recent outbreak of wasting syndrome. But I got distracted by other things and gave up on the stars, for now. I need to do some thinking about the best way to find tiny animals in a very complex 3-dimensional habitat.
I did spend quite a bit of time turning over rocks in tidepools. The most common critters I found were the usual suspects–porcelain crabs, limpets, snails, the odd sculpin or two, and chitons. One rock yielded a gold mine: five chitons of a species I didn’t recognize (which doesn’t mean I haven’t seen it before, just that I didn’t immediately know its name) that demonstrated a most interesting behavior.
I turned the rock over and watched as the chitons ran away from the exposed surface onto the other side. Yes, RAN. I’ve never seen a chiton do anything this fast. Chitons, for the most part, lead apparently inactive lives. When we do get to see them in their natural setting, at low tide, they are usually scrunched down hard on the rock waiting for the water to come back. Obviously they are much more active when covered with water, but we don’t get to see them then. In the lab, where they can be immersed all the time unless they crawl up the walls, they do wander around a bit; however, to see a chiton do much of anything requires time-lapse photography.
Don’t believe that a chiton can run? Well, get a load of this:
This is in real-time, not sped up. Watch the chiton push a limpet and the snail out of the way. Okay, I’ll grant that a limpet and a snail are not the strongest obstacles one could face when trying to flee from the light. But you can’t deny that this chiton seems to be feeling a sense of urgency.
This species, Stenoplax heathiana, spends its days buried in sand on the underside of rocks. It comes out to feed at night, not on algal scums as most chitons do, but on bits of algae that drift by and get caught between rocks. Apparently the chiton can be found exposed in the very early morning. I’m going to have to try finding some this spring when we get our morning low tides back. Anybody want to come with me?
