memory in cephalopods
I've heard the assertion that octopuses have short- and long-term memories several times in the past few days, mostly in discussions of the ethics of eating octopuses prompted by ethical questions raised about Paul, the famous German octopod. It's interesting to me what these people don't say - that they think that having a multiphasic memory process makes octopuses worth not eating (because, well, people have multiphasic memories, and you wouldn't eat them, would you?!? Sicko.) While I don't think that memory capacity of an animal is associated in an uncomplicated way with its ability to suffer or its moral status, it seems to me like a nonetheless interesting question. I'm almost sure that most of the people who use (read: copy and paste) this bit of information to support their beliefs have very little idea of what sort of research is behind it. Let's face it: developing a working knowledge of behavioral research on cephalopods is something that just isn't on most of the public's mind. In fact, until I began writing this blog, I had very little knowledge of the subject. I plan to set the record straight, so that internet users need never make an unfounded or unqualified statement about memory processes in cephalopods again (a lofty goal, huh?)
If you don't know octopus neuroanatomy very well (and who does?) you might want to check out the figures in this post. I'll be talking about the vertical and superior frontal lobes of the octopus brain, and I know it sometimes helps to be able to visualize things like that when you're reading about them. Just so that it's clear: the term "biphasic memory" means that the memory system in question has two discrete parts or processes (ie. short-term and long-term memory.) A monophasic memory would have only one process, so that memories would last for a certain amount of time and then fade similarly in all circumstances. A multiphasic memory system (which could be biphasic, triphasic, or more) is a general term to describe memory systems that are clearly more than monophasic, but are not completely characterized yet - and no memory system is. Now, on to the research!
J. Z. Young, that demigod of cephalopod neurobehavioral research, published one of the few papers I could find on this topic back in 1970, following up on his earlier work on the subject. In it, he investigated the development of short and long term memory in O. vulgaris (I assume - he doesn't actually mention what species he uses in this paper, but he almost always used O. vulgaris) as well as the role of two brain areas in memory, the median superior frontal lobe (MSF) and the vertical lobe (VL). To do so, he performed surgeries to remove one of these two areas of octopuses' brains and put them through a learning task. In this task, octopuses were trained to either attack a rectangle (rewarded with a piece of fish) or withhold attacking a crab (which was punished with electric shock.)
It turned out that octopuses whose vertical lobes had been removed were greatly impaired in learning to attack the rectangle. Young explains this by claiming that the vertical lobe is involved in short-term memory, and that the acquisition of stable behavior day-to-day was impaired because the animals without vertical lobes could not remember events long enough for the training to be effective. The animals without median superior frontal lobes, however, learned the task just fine, but were impaired in their long-term retention of it., suggesting that the MSF lobe might have some role in retaining learned information. Interestingly, Young also found (in other experiments) that removing the vertical lobe after a task was learned resulted in a greater retention of the task. These results suggest that the vertical lobe plays a role in the updating of memory stores, but is not absolutely essential for the recall of memories.
His results from the attack-withholding task were less clear, but they suggest that animals with lesions, especially those with vertical lobe lesions, were less consistent than intact animals in learning not to attack a crab after being shocked each time they attacked it.
Basically, Young argues (on the basis of this and some of his other experiments) that octopuses have a memory system that can be disrupted in more than one way; that is, it is possible to dissociate memory acquisition from long term retention, just like in vertebrates. For the most part, more current research has agreed with his position, as we'll see in this next paper.
Moving forward (past a lot of great research that I'll skip over for the sake of brevity) to 2008, Shomrat et al. used electrophysiological methods to test this hypothesis. Before we get into their methods, let's look a bit more closely at the system that we are talking about (this figure is from Shomrat et al. (2008)):
On the left is a sagittal slice of the supraoesophageal (over-the-oesophagus) mass of the octopus brain. On the right is a diagram of the memory system in question. Sensory information flows into the MSF from the arms and eyes before being sent along to the VL. The VL neurons in turn send out information encoding attack. It's been established that long-term potentiation (LTP) can occur in this area of the octopus brain, and this is a likely mechanism for the formation of memories in octopus (I blogged about this here - check it out if you need a little more background.)
The authors' procedure went as so: O. vulgaris who had already been trained to attack a white ball either had their MSF tract cut (at the dashed line in each image,) severing the sensory input to the vertical lobe, or this tract was stimulated, causing LTP at the synapses indicated in the figure. Shortly after the procedure, the animals were trained to avoid a red ball through electric shock. It was found that animals with severed MSF tracts were slower than controls to learn to withhold attack, while animals in whom LTP was induced were quicker. This is all well and good - it confirms what we already thought about the role of the vertical lobe in acquiring memories in the octopus. The really important result from this paper came when the authors tested the octopuses a day later. It was found that both MSF tract transection and LTP induction impaired recall after 24 hours. So even though stimulation of the MSF tract improved short-term memory (presumably by hyper-activating the memory system in the vertical lobe,) it impaired long-term memory. This suggests that these two processes are not identical; that is, that octopuses have discrete and dissociable short- and long-term memory circuits. This general finding has been replicated in cuttlefish (see my post on cuttlefish memory) and nautiluses (Crook and Basil, 2008).
Unfortunately, that's just about all that we know at this point: that cephalopods appear to have biphasic memories, meaning that the behavioral evidence of short-term memories can be dissociated from that of long-term memories. This is hardly (by itself) a basis on which we can imply any sort of consciousness or advanced cognitive capacity, as animal-rights supporters who mention this fact seem to imply.
In interpreting these results in the context of our knowledge of cephalopods as a whole, we should keep in mind what is meant by short- and long-term memory in humans. Short-term memory is what happens when newly learned information is bouncing around the cortex somewhere, being continually processed but not permanently encoded somewhere. These memories will disappear if they are not rehearsed (or otherwise actively retained). Long-term memory has been (relatively permanently) encoded into neural circuits, so that it can be retrieved after periods when it has not been actively processed in short-term (or working) memory circuits. These processes have been studied intensely in humans, and can be precisely because we have a complex cognitive system build around them (or on top of or parallel to them, depending on who you ask) that we can access. As of yet, we don't have the experimental techniques to assess exactly how "human-like" or "vertebrate-like" cephalopod memory systems are, because we can't study them in nearly as much detail as language-based and other cognitive tasks allow us to in humans. Thus, making any strong conclusions about the nature of cephalopod memory other than that it appears to be multiphasic (with no implied "and-so-cephalopods-are-smart-like-people") is untenable.
Lastly, I find it frustrating that animal rights activists use our (very primative) knowledge of cephalopod memory systems to try to support their position that eating cephalopods is wrong. Not only is it an inconclusive (what does memory have to do with suffering and morality?) and nonspecific argument (did anybody think that ungulates, swine and birds don't have complex memory systems?), but it misses some of the big points that the animal rights movement has taught us. First of all, it implies that cephalopods are somehow special because they are intelligent and human-like. However, having compassion for animals explicitly demands that we not judge their worth by analogy to our own abilities - this has proved to be an attitude that encourages cruelty to animals simply because we are ignorant of them and their behavioral and cognitive capacities. If we didn't know about cephalopod memory systems, would they still be worth defending from fishing and consumption as food? Hopefully, the answer is yes - so why try to use this (admittedly inadequate) argument now that we conveniently have information that appeals to one's emotional predispositions? I find this to be irresponsible and counter-productive, as it diminshes the credibility of other, more valid arguments against the consumption of cephalopods (or any animal, for that matter) that animal rights activists might use.
Sorry if this was a bit heavy on editorial material. Being very concerned about animal welfare myself, I get annoyed when people make the cause look stupid by saying things that are ill-informed, ill-reasoned, or just plain wrong. Although I wish that people would stop killing cephalopods for food, spinning information to try to get people to agree with a point is dishonest, and at best a very poor strategy for debate, as there's bound to be at least one attentive person on the other side who will point out that you're not being true to the facts - and nobody will listen to you after that.
Thanks for reading!
SHOMRAT, T., ZARRELLA, I., FIORITO, G., & HOCHNER, B. (2008). The Octopus Vertical Lobe Modulates Short-Term Learning Rate and Uses LTP to Acquire Long-Term Memory Current Biology, 18 (5), 337-342 DOI: 10.1016/j.cub.2008.01.056
J. Z. Young (1970). SHORT AND LONG MEMORIES IN OCTOPUS AND THE INFLUENCE OF THE VERTICAL LOBE SYSTEM Journal of Experimental Biology (52), 385-393
Crook, R., & Basil, J. (2008). A biphasic memory curve in the chambered nautilus, Nautilus pompilius L. (Cephalopoda: Nautiloidea) Journal of Experimental Biology, 211 (12), 1992-1998 DOI: 10.1242/jeb.018531
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