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Tuesday, October 27, 2015
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Cephalopod Links
First things first: a few recent cephalopods in popular news media:
Paul, the German octopus, successfully predicted yet another World Cup match. An aquarium in Oberhausen, Germany had its animals place their bets on World Cup matches as a promotion, and the world discovered that Paul is the luckiest octopus to ever live.
In Australia, three men are arrested for stealing cuttlefish from a protected area of ocean. The town police consider selling the catch as food - failing this, they will bury it. I couldn't make this up.
In Oregon, researchers try to recruit fishermen to help them understand Humboldt squid population dynamics off the Oregon coast.
And some topical blog posts:
Marine Biologist William Gilley updates us on the progress of an expedition to study Humboldt squid in the Gulf of California.
A recipe for char-grilled baby octopus salad at Food Stories (this makes me sad, oh so sad, but I had to post it for the weirdness.)
Richard Ross gets very excited about a pair of Metasepia pfefferi getting it on (with video!) I have to say, of all the animal mating rituals one could watch, cuttlefish are pretty easy on the eyes. In all seriousness, though, culturing cephalopods is notoriously difficult, so this is probably a huge deal.
The Carnival of Evolution #25 is out at Culturing Science - take a moment learn about empathy in crows and all the other amazing things that the evolutionarily-minded blogosphere finds fit to write about. You'll be glad you did.
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Know on systems sensual octopus
Know on systems sensual octopus
This will be a quick one - I'll get back to the meat of my series on octopus sensory systems soon, but I wanted to write a post on this article because it struck me as cool (although it has a sort of sensational title.)
The article I'm talking about is Octopuses (Enteroctopus dofleini) Recognize Individual Humans (2010) by Anderson et al. in the Journal of Applied Animal Welfare Science.
The authors used an apparently elegant experimental design to test whether octopuses can tell people from one another across a long period of time - specifically, this is operationally defined as meaning that they could learn an association between a person's features and a good or bad stimulus. The experiment was conducted thus: eight octopuses were captured and habituated to their aquaria. Then, for 2 weeks, the octopuses had daily interaction with two people, one of whom fed them and one of whom (I'm not joking) poked them with a "bristly stick" (more specifically, "a length of PVC pipe with one end wrapped in Astroturf.") Then, the octopuses were tested to see if they reacted differently to the two individuals - presumably, if they remember who is who, they should show anticipatory behaviors related to eating or defensive behaviors in response to the appropriate person.
To get a better feel for the task, here are the experimenters, shown in an image taken from the octopus's point of view:
My problem with this experiment is that the term "individual" is usually used in cognitive research to mean some entity who is known to persist despite changes in their appearence in one specific sensory modality. When we get a haircut, our friends (and, usually, our pet dogs and cats) still recognize us - thus, we are individuals to them. However, if the visual stimulus of the two keepers didn't change from day to day (and they took pains to make sure that it didn't,) then this seems like little more than a complex visual discrimination task. It seems, judging from this image, that it would be pretty easy for an octopus to learn an association between, say, a shiny bald head and being jabbed with a stick, regardless of any ability she might have to recognize "individuals" in the cognitive sense. In any case, we are still a ways away from knowing whether octopuses can recognize individuals, and not just their constant visual features. With this in mind, let's consider their results.
It turns out that the octopuses learned to move away from the irritator and towards the feeder within two weeks. In addition, the octopuses showed fewer defensive coloration responses to the feeders than to the irritators, as well as changes in their respiration rate and the orientation of their bodies relative to the people. In sum, it looks like (in this test, at least) the octopuses succeeded in learning basic traits about the people interacting with them. I don't think that the title of the paper is fully supported, however - it's hard to make the case that this single study proves that octopuses can identify individuals in any sort of robust way.
This paper is pretty solid (besides its unfounded title,) although it begs a few questions:
1. How fine of a discrimination can octopuses make? Would they treat two bald men of similar stature the same? What if the subjects wear different clothes? How is this piece of research fundamentally different from Wells' experiments using simple visual cues? These are all important questions if we're actually going to claim that octopuses can identify "individuals" as opposed to simple visual stimuli.
2. What does this mean functionally to the octopus in the wild? Is this sort of ability actually used to identify predators and prey items? Do octopuses remember individuals of any species in the wild? Unfortunately, there is not much literature on the development of behavior in the octopus, so we can't know how much of octopus behavior is "instinct" and how much of it is based on learning (like that shown in this study.)
3. How does this generalize to other species of octopus? This study used Enteroctopus dofleini, the giant pacific octopus, because it is often kept in public aquaria. However, practically the whole body of research on octopus learning and vision has been done using O. vulgaris and, to a lesser extent, O. cyanea. We know that cephalopods have a pretty wide diversity of life-styles, so it seems important to me to know how these behaviors occur in different species if findings like this are going to be relevent to the rest of cephalopod research.
If nothing else, this study keeps alive my childish hope that Twister, the resident E. dofleini at the Niagara Falls Aquarium (which I visit almost weekly these days) will someday get to know me, if only in the most basic way.
Anyways, I hope this has been as fun for you as it was for me. Thanks for reading!
Anderson, R., Mather, J., Monette, M., & Zimsen, S. (2010). Octopuses (Enteroctopus dofleini) Recognize Individual Humans Journal of Applied Animal Welfare Science, 13 (3), 261-272 DOI: 10.1080/10888705.2010.483892
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Poisonous octopus in Antarctica
Poisonous octopus in Antarctica
In my recent quest to find new, cutting-edge research on cephalopods, I've come across some neat stuff (check out this post on the perception of polarized light by cuttlefish - it's one of my favorite new cephalopod research topics!) The study I'll review here is outside of my field of relative expertise, but it's so neat and so new that I couldn't resist writing about it. It's good to step out of one's comfort zone every once in a while, right?
An international team of researchers hailing from Norway, Australia, and Germany has published a study on the venom of Antarctic octopods (more accurately, it is being published, though it hasn't hit the presses yet.) The team investigated the biochemical properties of extracts from the salivary gland of four Antarctica octopus species and wrote up their results in Venom on ice: First insights into Antarctic octopus venoms (2010).
Here is their image of the posterior salivary glands of an octopus, from which the authors collected all of their specimens:
These glands produce a variety of compounds, notably venom and digestive enzymes. The venom of temperate-water octopuses has been studied in the past. Never before, however, has venom been studied in an octopus that lives in below-freezing temperatures, conditions under which the enzymes in most venoms work very poorly if at all. To begin to understand the role of venoms in the lives of Antarctic octopuses, the team collected and tested venom from four octopus species collected off the Antarctic shore: Adelieledone polymorpha, Megaleledone setebos, Pareledone aequipapillae, and Pareledone turqueti. Here are images of some of their specimens:
Cute, aren't they? Octopods always are! Anyways, back to the biochemical assays.
First, the authors tested the extracts for alkaline phosphatase (ALP) activity. ALP is an enzyme that is in spider and snake venom that is thought to help immobilize prey items. Second, they tested for Acetylcholinesterase (AChE) activity. AChE breaks down acetylcholine, a neurotransmitter, potentially acting as a toxin by disrupting neuromuscular function. Third, the extracts were tested for general proteolytic activity using casein. Fourth, an assay for secreted phospholipase A2 (sPLA2) was performed. sPLA2 is found in cone snail and snake venome, and contributes to the effects of venoms in a variety of ways. Finally, the researches assessed whether the venoms showed haemolytic activity, which is a common marker of the general toxic activity of venoms. Taken together, these results should begin to characterize the putative venom of each octopus species. After all of this, the researchers reviewed what is known about the morphology of the mouthparts of the octopuses, as well as their feeding habits, and tried to relate these to their biochemical findings.
Whew.
So, after all of that, what did they find? Here are their results(takes another deep breath):
Venom from all of the species had some ALP activity. Interestingly, however, when ALP activity was tested at 0 Celsius and at 37 Celsius, venom from 3 species of octopus (A. polymorpha, M. setebos, and P. turqueti) had higher ALP activity at the lower temperature! This is a significant finding because it suggests some sort of modification of the proteins responsible for this activity to function optimally at a lower temperature. This lends some weight to the theory that the use of venom has been important enough to the survival of Antarctic octopus species that they have evolved enzymes to work under conditions where most enzymatic toxins would not. In the other tests, an essentially similar pattern of results were found, except for the AChA activity assay. Little AChA activity was found in any of the species, although the results of the assay were poor enough (that is, inconsistent) that they were not included in the paper. Interestingly enough, although all of the species had a few potentially functional toxins in their venom, most of them showed only weak haemolytic activity. Only one extract (from P. turqueti) showed strong haemolytic activity.
The relation of venom activity to morphology and diet that the authors attempted to point out appears to be weak (or at least difficult to point out given their sample,) as it is mentioned that few clear venom-related adaptations in diet or anatomy were present in these octopus species. A. polymorpha is noted to have a very large salival gland and a narrow beak, which the authors suggest might be an adaptation associated with the use of venom as a primary means of catching prey (as opposed to having powerful jaws to physically overpower the prey.) This species feeds mostly on amphipods and polychaete worms, and so it's unclear why it would rely on venom to subdue such (relatively) easy going prey instead of retaining a more varied diet. In any case, though, this is one of those papers that, being exploratory, raises many more questions than it answers - that's the kind I like!
What I find most interesting about this work is that it begs questions about the evolution of octopus venom. How quickly could the octopus populations move into cold water? Was this limited by the evolution of venom enzymes, or did that evolution occur after some quicker relocation of the species which left their warm-water-adapted enzymes useless? Did A. polymorpha's ancestors have a specialized diet before they became Antarctic, or is that only a successful feeding strategy in the Antarctic environment? The world may never know (although I hope we do, someday!)
Thanks for reading!
Undheim, E.A.B., et al. (2010). Venom on Ice: First insights into Antarctic octopus venoms Toxicon
Visual ability to see the squid
Visual ability to see the squid
I thought I'd mix things up a little bit and take a look at some research on the sensory abilities of cuttlefish. Specifically, I'd like to take a look at an aspect of cuttlefish vision that has shown up in the literature recently (it's actually one of the few threads of cuttlefish research that seems to be active at the moment - the other ones I've noticed are memory and fishery ecology and management): the ability of cuttlefish to perceive polarized light. Polarized light is composed of photons that are all oscillating in the same plane - we cannot sense the polarization of light, but it seems to play some role in the lives of cephalopods and some other animals. For more info on polarized light, check out this explanation of polarization.
It has been known that cephalopods can respond to polarized light for some time - Wells did the work showing that octopuses can detect polarized light in the 1960's, and it's been studied in fits and spurts since then. In the late 1990's and early 2000's (from what I can tell,) it became a relatively hot topic among researchers who study animal communication, because it appeared as if cuttlefish might be able to use polarized light for some sort of intraspecific communication. A good though somewhat dated review of the topic is Shashar et al's Polarization Vision in Cuttlefish - a Concealed Communication Channel? (1996).
How can cephalopods see polarized light? It turns out that their photoreceptors are orientated at a variety of angles, so that incoming light will cause the most stimulation in photoreceptors that are oriented the "right" way. In unpolarized light, all of the cells would be pretty much equally stimulated - nothing unusual happens here. Upon being hit by polarized light, though, a specific population of retinal cells (those that are oriented in the proper direction) will be activated, and the animal will be able to see the polarization of light.
This is an image of cuttlefish (S. officinalis) photoreceptors (From Shashar et al 1996.) The lines are folds in the photoreceptor cells called microvilli. Notice how the two adjacent cells have microvilli at a right angle to each other - this is what allows cephalopods to see the difference between polarized and non-polarized light.
Detecting polarization can help a creature in a lot of ways. In a basic sense, it almost always helps an animal (especially one who, like the cuttlefish, is both a predator and a prey item) to have as much information about the environment. If sensing polarization allows the cuttlefish to know more about its environment at any given time, it's already a huge advantage. In fact, it has been shown that the perception of polarized light probably helps cuttlefish to catch certain prey that is difficult to see otherwise (see Shashar et al 2000.) But I mentioned the possibility of communication through polarized light - how does that work?
It turns out that iridophores, organs in the skin of cephalopods that reflect light, polarize that light to some extent. The anatomy of iridophores is such that they preferentially reflect light polarized in a certain plane. It is known that cephalopods, especially cuttlefish, have wonderful neural control over the pigment organs in their skin, which allows them to display such a dazzling array of colors and patterns. Cuttlefish might be able to manipulate the polarizing properties of their iridophores, adding another layer of complexity to their body patterns. Importantly, however, this would be a type of display that not everybody in the sea could perceive. Shashar's theory is that cuttlefish might use polarized light as a type of social signal, while still being able to maintain the camoflauge which is key to avoiding being eaten.
Shashar and friends did a few experiments to test this hypothesis: first, they observed cuttlefish during a variety of behaviors, and found that the polarization of light being reflected from the cuttlefish's arms varied with different behaviors in much the same way as their patterns of coloration. Polarized light is reflected from stripes on the arms and the area around the eye, as seen in this image from a review on the use of polarized light by cuttlefish by Mathger et al (2009):
The top image is a cuttlefish as seen by the human eye. The bottom image has been given false color, so that areas which reflect polarized light show up as green. On an unrelated note, cuttlefish sure are cute.
In addition to discovering the patterns of reflection of polarized light by cuttlefish skin, the authors found that cuttlefish respond differently to their own reflections when they view them through a filter that screens out polarized light. Specifically, they found that cuttlefish responded less noticibly to the disrupted image. While the authors declare that these findings are "fully consistent with the hypothesis that cuttlefish use controllable polarization patterns for intraspecific communication," they are also consistent with the more parsimonious explanation that cuttlefish don't respond to any stimulus made of non-polarized light as strongly as they do when it is at least partially polarized. While the theoretical argument presented in this paper is interesting, I think it's a bit too eager for what the data show.
Fast-forward to 2004: Boal et al. published a study called Behavioral evidence for intraspecific signalling with achromatic and polarized light by cuttlefish. In this study, they exposed cuttlefish (S. officinalis) to conspecifics (that is, other cuttlefish) through either a clear or a polarized light-blocking barrier. They found that only females responded differentially to conspecifics behind the polarization-distorting barrier, not responding to them at all (cuttlefish confronting each other unexpectedly often show some sort of postural and color change.) This was the only significant result that they found, and it is ambiguous in its interpretation. Again, it might simply be that a non-polarized stimulus is not very interesting to an animal who is used to seeing a world of polarized light.
So, do cuttlefish use polarized light to communicate? I'm not convinced. It seems as if everybody's hoping that it's true, but there's not any good data showing it to be so. I can't sum it up any better than Mathger et al. did in their 2009 review:
The fact that cephalopods can detect polarized light
and can also produce changeable polarized light
patterns in their skin begs the question whether
cephalopods communicate using polarized light signals.
The likely answer is that they do. Unfortunately, we
have little evidence to support this statement.
Thanks for reading!
Shashar N, Rutledge P, & Cronin T (1996). Polarization vision in cuttlefish in a concealed communication channel? The Journal of experimental biology, 199 (Pt 9), 2077-84 PMID: 9319987
Mathger, L., Shashar, N., & Hanlon, R. (2009). Do cephalopods communicate using polarized light reflections from their skin? Journal of Experimental Biology, 212 (14), 2133-2140 DOI: 10.1242/jeb.020800
Boal, J., Shashar, N., Grable, M., Vaughan, K., Loew, E., & Hanlon, R. (2004). Behavioral evidence for intraspecific signaling with achromatic and polarized light by cuttlefish (Mollusca: Cephalopoda) Behaviour, 141 (7), 837-861 DOI: 10.1163/1568539042265662
Shashar N, Hagan R, Boal JG, & Hanlon RT (2000). Cuttlefish use polarization sensitivity in predation on silvery fish. Vision research, 40 (1), 71-5 PMID: 10768043
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