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Can Whales and Dolphins Make Mental Maps?

The views expressed are those of the author and are not necessarily those of Scientific American.

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The brains of cetaceans—dolphins and whales—differ from those of other mammals in a number of ways, but one of the most striking differences is the size of the hippocampus.  As a general rule, the larger the size of a mammal’s brain, the smaller the fraction of it that the hippocampus occupies, so dolphins and whales would be expected to have a small hippocampus in any case. But the cetacean hippocampus isn’t just small; it is so tiny that it barely exists.

The relative size of the cetacean hippocampus was recently quantified by a group of researchers led by Paul Manger (Patzke et al, 2013). They examined data on the size of the hippocampus in several hundred species of mammals, including several species of whales and dolphins. They found that a plot of hippocampus volume versus total brain volume yields points that are almost all clustered tightly around a smooth curve—but the points for cetaceans are outliers. Their hippocampal volumes are only 8 percent to 20 percent of what would be expected on the basis of total brain size. No other type of mammal comes close to matching that. Even the hippopotamus—the nearest living relative of cetaceans—has a hippocampus size close to what the main trend line predicts

Log-log plot of hippocampal volume versus total brain volume, for 375 species of mammals from a wide variety of groups. Used with permission from Patzke et al, 2013.

Manger’s group also found another striking difference between the hippocampus of cetaceans and other mammals. The hippocampus is one of only two brain areas that are known to show neurogenesis (creation of new neurons) in adult mammals (the other is the olfactory bulb). Manger and his colleagues examined the hippocampus of 71 species of mammals and found evidence for adult neurogenesis in all of them except the ones from cetaceans.

Manger has long been known as a skeptic about the intelligence of dolphins and whales, and he interprets the data as additional evidence that their brains are not as sophisticated as the brains of other mammals. It is interesting, though, to think about the findings in light of current theories of hippocampal function. At a general level, the small size of the cetacean hippocampus and the absence of adult neurogenesis both suggest the same conclusion: whatever function the hippocampus performs, dolphins and whales don’t have as much need for it as other mammals.

But what function is that, exactly? Executive summary of the answer: we can guess but we don’t know. There are two main functions that theorists associate with the hippocampus, and both involve memory. The first is long-term episodic memory; the second is spatial memory. I will consider each in turn.

Credit: Gregory "Greg" Smith via Flickr

In humans, destruction of the hippocampus causes inability to form new memories and major loss of memories from the recent past, but memories from the distant past, such as childhood, are usually intact. A patient with this type of amnesia can meet with the same doctor daily for years without ever learning the doctor’s name or even knowing that he had met this person before.

It is important to clarify that hippocampal amnesia does not affect all types of memory. Procedural learning (the ability to learn new tasks) and working memory (the ability to hold information in mind briefly) remain intact. The signature of hippocampal amnesia is an absence of conscious awareness of having experienced an event, not an absence of memory in general.

So what about dolphins and whales? Is it possible that their diminutive hippocampus means that they lack conscious memory of past events? Unfortunately there is not enough data to answer the question. There have been several experimental studies of memory in dolphins, but not the right sort of memory.

There is one sort of memory that dolphins clearly possess to a high degree (Herman, 2010). A dolphin can be instructed, using a gesture-language, to perform a series of actions, and then a short time later it can be instructed to “repeat” them. Dolphins are quite good at that. But the delay between the first performance and the repetition is generally only a matter of seconds, so this has to be classified as working memory rather than episodic memory.

Credit: Jay Ebberly via Flickr

As far as episodic memory is concerned, the most relevant information comes from a study last year that found that dolphins are capable of recognizing the vocal calls of other dolphins they have previously known, even after an absence of decades (Bruck, 2013). If recognizing a vocal call is equivalent to learning a person’s name, then this ability would be unexpected in hippocampal amnesia. But it would be dangerous to draw strong conclusions from a single piece of information such as this.

In summary, dolphin episodic memory is a fertile area for future experimental work, and we can’t yet predict how it will turn out.

What about spatial memory?

One of the most popular theories of hippocampal function over the years has been the “cognitive map hypothesis” (O’Keefe and Nadel, 1978), which says that the hippocampus plays a central role in the formation of mental maps, allowing the brain to represent the layout of an environment and the location of objects and features.

It is interesting to note that cetaceans in their natural environment probably have less need for cognitive maps than any other type of mammals. They are the only mammals that spend most of their lives at the surface of the open sea. On the open sea there is no meaningful concept of a “place”. Spatial relationships are very important, of course, but they are relationships between pairs of objects, not between an object and a place in a stable environment. (I am talking about small-scale maps here. Dolphins and whales clearly would benefit from being able to learn the general features of a large area of space—the layout of coastlines, the locations of islands, reefs and river outlets, etc. But there are reasons to believe that large-scale maps of that type rely on different neural mechanisms from small-scale cognitive maps.)

Credit: Réunion Underwater via Flickr

The question, then, is whether dolphins and whales are able to form mental maps of an environment and remember the locations of objects. It should be reasonably straightforward to test that experimentally, because even though the natural environment of free dolphins is unstructured, captive dolphins spend their lives in tanks with limited size and fixed features. It makes sense to ask how well they understand the spatial layout of the tanks they live in. Unfortunately here too experimental data is almost entirely lacking. Scientists who study spatial cognition have worked out a number of systematic ways of testing spatial memory, but none of them have been attempted using dolphins.

Practically the only relevant information comes from a study carried out by Kelly Jaakkola and her colleagues (Jaakkola et al, 2010). They wanted to know the extent to which a dolphin could maintain a memory of the location of a hidden object. Their procedure involved a small stuffed toy and three buckets spaced a foot apart at the edge of the dolphin’s pool. The toy was either (1) placed in one of the buckets in full view of the dolphin; (2) placed in one of the buckets and then moved to a different bucket; (3) placed in a container and then invisibly transferred to one of the buckets; (4) placed in a bucket, after which that bucket was transposed with another one. The general finding was that the dolphins succeeded in conditions 1 and 2 at above-chance levels, but not in conditions 3 and 4. Even where they succeeded, performance was erratic and a great deal of careful training was required. The authors concluded that dolphins have only a limited ability to mentally track the locations of hidden objects—roughly as good as dogs, but not as good as great apes.

Beyond that, the only information available is anecdotal. Louis Herman and Paul Forestell carried out an experiment in the 1980s in which they asked dolphins to report the presence or absence of named objects in their pool (Herman and Forestell, 1986). In many cases the dolphins would take time to explore the pool before responding. In some cases the dolphins would respond in a way that suggested awareness, including awareness of what part of the pool the object was located in, but that aspect was not examined formally.

The bottom line is that we really have no solid information at this point about whether dolphins can form cognitive maps. And the cumulative bottom line is that we can speculate about whether the tiny hippocampus of dolphins and whales brings about a weakness in episodic memory or spatial memory, but at this point we just simply don’t know the answer.

William Skaggs About the Author: William Skaggs is a neuroscientist whose experimental work has focused on the role of the hippocampus in learning, memory, and spatial navigation, but he is interested in several other areas of science as well, especially the study of consciousness. He has ambitions to be a science writer, and has contributed extensively to Wikipedia under the name "Looie496", mainly by writing articles about the nervous system. Follow on Twitter @weskaggs.

The views expressed are those of the author and are not necessarily those of Scientific American.

Comments 9 Comments

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  1. 1. tuned 11:17 am 04/25/2014

    Any intelligence is subject to various definitions and viewpoints.
    I saw a video of a white blood cell chasing down a bacteria.
    I was amazing. It chased it around other cells, etc. until it intercepted the bacteria and killed it.
    How did it differentiate them all?
    How did it know where itself was? How did it know where the bacteria was when the bacteria went around another cell?
    Why did it care?
    It looked like anything but pinball arcade action-reaction. Fascinating.

    Lastly, is there anything about the hippocampus matter that can not be accomplished in the other (mostly grey) brain matter if “nature says” it need be?

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  2. 2. BillSkaggs 4:11 pm 04/25/2014

    Hi, “tuned”. As I tried to say in the blog post, if the human hippocampus is destroyed, the result is amnesia. So in humans at least, the hippocampus is necessary for properly functioning memory. Perhaps in principle other parts of the brain would be capable of carrying out that function, but in actual fact they don’t.

    Regards, Bill

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  3. 3. KS14143501 3:44 am 05/1/2014

    You mentioned that they used mainly captive animals. Wouldn’t it be more prudent to test these hypotheses on wild animals rather than their captive counter-parts.

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  4. 4. BillSkaggs 9:01 am 05/1/2014

    Hi, “KS”. There are a couple of reasons why captive dolphins are used. First, the logistics of working with free-ranging dolphins would be very challenging. Second, virtually all experimental work on dolphin psychology has been done by people who work as dolphin trainers at places like Sea World. You’re right that this raises issues, but it’s hard to see any practical way of avoiding them, at least without major increases in funding for this type of research.

    Regards, Bill

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  5. 5. davantlag 1:08 pm 05/1/2014

    I am not totally sure, despite tha fact that this habit apparently makes everything easier, that we should use our brain model as a “paramount” or a “rosetta stone” for any other on the Earth. Maybe in the immensity of the oceans whales and dolphins’s brains are wired in a different way since they need a special adaptation to their environment. We should remember that dolphins can split their brain in two, allowing an hemisphere sleep meanwhile the opposite part is still wide awake and leading. How can we compare?

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  6. 6. KS14143501 2:23 pm 05/1/2014

    I agree with “davantlag”. These animals have adapted in incredible ways. We know very little of our own brain so why compare everything else to it? Is it even possible to map a brain of an animal like a dolphin or a whale?

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  7. 7. KS14143501 3:51 am 05/2/2014

    Do whales and dolphins not return to the same spot to mate and reproduce? if so isn’t that an indication that they can form mental maps?

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  8. 8. BillSkaggs 10:19 am 05/5/2014

    Hi devantlag, I agree with you, but to me the fact that something might be hard to understand is an incentive to study it.

    Hi “KS”, I don’t know anything about that phenomenon, but even if they do return to a “spot”, it surely must be a spot in the sense of a region rather than a point, and that wouldn’t contradict anything I wrote.

    Best regards, Bill

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  9. 9. enrico 10:11 am 02/21/2015

    …Yet gradually, over a period of two to three decades, despite its glossy pages, added color, web links and other bells and whistles, the magazine has become a shadow of itself. It is a story that is not too different from what has happened to the content of most high school science courses and textbooks….

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