What is Animal Cognition?

Lecture 1

“Housekeeping”

Cognition, broadly defined, means the processing of

information in order to make informed choices…

but when animals are concerned, the definition

becomes muddied

Views of animal cognition seem to swing, pendulum-style, from one extreme

view to another….

Mindless robot

Furred or feathered

human

Although we can could discuss humans’ views of

animals in numerous civilizations throughout

recorded history,

we’ll concentrate on Western, fairly modern

civilization, if only to keep this lecture to within a

reasonable length

On the mindless robot side, we have proponents like Descartes

and the mid-20th century behaviorists…..

On the furred, feathered human side we have proponents like

Romanes and Griffin.

I hope at the end of the class, you’ll come to

believe that the answer lies somewhere in the

middle…

In his Meditations in the 1600s, Rene Descartes argued that

animals are purely physical entities,

lack mental and spiritual substance,

can’t reason, think, feel pain or suffer,

are machines with no consciousness

He surmised that they might have some fleeting responses to stimuli,

but argued that they definitely could not think because they

couldn’t use or didn’t have language demonstrated behavior that was not all that adaptable

reacted mostly instinctually

Now, Descartes was not the only one to present this

stance, but his writings were widely accepted and fit in with the general view of animals as

creatures that existed only for the benefit of humans:

as sources of food, beasts of burden

And it’s much more difficult to justify such a

stance

if one believes that animals are sentient beings,

more similar to, than different from, humans

Although, as we will see, no animal exhibits behavior that is identical to

that of humans

suggesting a continuum

so as to totally contradict Descartes,

many animals exhibit behavior patterns with many elements that

approach that of humans,

In the 1800s, Romanes, on the heels of the

publications of Darwin,

began to address the inconsistencies in

Descartes’ position

In 1871, Darwin stated: “the difference in mind between man and the higher animals…is one

of degree, not of kind”

To some extent, Darwin, but mostly Romanes,

went about collecting a huge number of anecdotes to support

this idea

Romanes classified these anecdotes by phyla and species

The idea was to find a precise phyletic point at which a particular

“new” mental ability emerged

in order to establish a continuum from single-celled

organisms to humans….

The upshot were great “chains of reason”, with humans at a

pinnacle;

the big problem was that Romanes’ material was simply

anecdotal,

An example:

depending on human recollections and lacking any

level of control

Another time I found a very few of them passing along at intervals. I confined one of these under a piece of clay at a little distance from the line, with his head projecting. Several ants passed it, but at least one discovered it and tried to pull it out, but could not. It immediately set off at a great rate, and I thought it had deserted its comrade, but it had only gone for assistance, for in a short time about a dozen ants come hurrying up, evidently fully informed of the circumstances of the case, for they made directly for their imprisoned comrade and soon set him free. I do not see how this action could be instinctive. It was sympathetic help, such as man only among the higher mammalia shows. The excitement and ardour with which they carried on their unflagging exertions for the rescue of their comrade could not have been greater if they had been human beings.

         This observation seems unequivocal as proving fellow- feeling and sympathy, so far as we can trace any analogy between the emotions of the higher animals and those of insects.

On ants:

This material, as written, probably seems totally off-the-wall and

unbelievable….

And it was just these kinds of reports that really acted to

discredit Romanes—for decades— as a serious scientist

Moreover, neurobiology seemed at first to be against Romanes

Researchers argued that certain brain areas—

And, creatures such as reptiles and birds lacked this brain area

specifically those in what was called the cerebral cortex—

were necessary for intelligent behavior

Songbird Human

And even in other mammals, including our nearest relatives, the apes,

even if, as much later researchers argued,

the area was considerably smaller….

one took into account brain-body ratios

From Jerison, 1955

But, before we go on….

So it seemed as though the term “animal intelligence” or

“animal cognition”

Let me get jump ahead chronologically in terms of

brain data …

was an oxymoron…

First, papers that came out just about two years ago show

that birds, at least,

but that for some species,

not only have bits of brain that are equivalent to human cerebral

cortex,

this area has about the same brain/body ratio as in primates

Fig. 10. Scatterplot using independent contrasts of primate and psittaciform data, of log-transformed telencephalic volumes (mm3) against log-transformed body mass (g) contrasts; Psittaciforms are indicated by the gray circles and the primates are indicated by the open circles.

And, as we will see later in the semester,

those birds with such large areas (the crow and parrot

family)

perform at about the level of young humans

Second, an article in the January 12th, 2006 issue of the journal Nature

reports evidence for teaching in ants:

They performed less efficiently than they would outside the “classroom”

Their pupils learned faster than they would have learned by themselves

Feedback existed between teacher and pupil

[go to desktop]

So, although Romanes likely over-interpreted what he did find,

But back at the turn of the last century,

some grain of truth likely existed in his material

Romanes’ material was thought to be so outre

that the pendulum swung wildly in the opposite direction….

Enter Morgan, who, in reaction to Romanes’ “mentalistic” approach,

In no case may we interpret an action as the outcome of the exercise of a higher psychical faculty, if it can be interpreted as the outcome of the

exercise of one which stands lower in the psychological scale (1894).

formed his famous canon:

Or, in its revised version of 1903:

In no case is an animal activity to be interpreted in terms of higher psychological processes, if it can be fairly interpreted in terms of processes which stand lower in

the scale of psychological evolution and development.

Add the comments of Thorndike (1898):

“… anecdotes give really the abnormal or super-normal psychology of animals.”

Scientists “…have looked for the intelligent and unusual and neglected the stupid and normal.”

“Only a single case is studied, and so the results are not necessarily true of the type; the observation is not repeated, nor are the conditions perfectly regulated; the previous history of the animal in question is not known.”

And:

“Such observations may tell us, if the observer is perfectly reliable, that a certain thing takes place, but they cannot assure us that it will take place universally among the animals of that species, or universally with the same animal.” “Nor can the influence of previous experience be estimated.”

Basically, the argument was that if a behavior could be

interpreted as a simple response to a given stimulus,

THAT was how it should be viewed.

an instinctual reaction,

or anything that did not require any level of information processing,

Morgan’s canon led to a very different approach to animal studies….

The good part was that careful experimental design became an

important criterion

Again, from Thorndike:

“To remedy these defects experiment must be substituted for observation and the collection of anecdotes.”

Thorndike argued that:

“You can repeat the conditions at will, so as to see whether or not the animal's behavior is due to mere coincidence.”

“A number of animals can be subjected to the same test, so as to attain typical results.”

“The animal may be put in situations where its conduct is especially instructive.”

Whether the next part was good or bad depends on

one’s point of view…

The idea that there was some kind of continuum of intelligence

began to wane

Instead, researchers became concerned with discovering

general mechanisms and laws of learning

that applied across all phyla

Although researchers expected to find quantitative differences in learning

across species,

they believed that the underlying principles would be the same.

And, critically, all such experiments were to be carried out in a sterile lab,

where all conditions could be controlled.

By isolating the animal,

one would avoid all extraneous input

and be able to determine what was left

—the basic ability—

with respect to intelligence

Of course, such a stance did not appreciate that the

absence of social stimuli

of the environment or of other individuals

was in and of itself a specific—and unnatural—situation for most

animals

Models of behavior were now to be based on associationist principles:

Simple laws were formulated to explain how external sensory input caused

observable behavior

Laws were few and were universal

(i.e., standard “stimulus-response” interpretations)

Thus there was no need to examine “cognitive” skills,

which, in part, included learning, remembering,

problem-solving, rule and concept formation,

perception and recognition

And one didn’t need to study a variety of species,

And only the receipt of reward or punishment controlled behavior

because none would respond any differently from that of a pigeon

or a rat

So, an animal would see an environmental stimulus, such as a green-lit button in one location, and at some point in its random

behavior it would come across another green button; if it pressed the button it got some

rewardA weak association thus formed between “green” and reward; if the animal hit a

nearby red button instead, nothing happened, or it got a shock…and this latter behavior would thereby be ‘extinguished’

The next time the rat saw the buttons, hitting green was more likely because of the reward

At issue was not a memory for reward:

Although some researchers did argue for memory, most didn’t care how the

rat chose to behave,

And researchers argued that any complex behavior could be broken

down into a series of such simple S/R laws

but rather only the extent to which it DID behave..

These researchers, known as behaviorists, did have some initial

success.

They could explain how animals learned via trial-and-error on a

number of laboratory tasks

(things like maze running or adapting when the experimenter

reversed the correct and incorrect choices)

And they had some success on ‘learning sets’ (e.g.,

Harlow), in which animals learned one task on one set of exemplars (e.g., to

choose the odd object among three)

and then responded more quickly when given the same task with different exemplars.

Ditto for Thorndike’s puzzle boxes,

which were used to show not only trial-and-error learning,

but also species differences in such abilities

Interestingly, these boxes showed that dogs and cats were

particularly incompetent….

But from the perspective of a dog or a cat,

The animal never was shown the correct response.

what did pulling a string have to do with opening a door?

And, in nature, animals rarely can rely on trial-and-error learning….

They get only one chance to avoid a predator, and only a limited amount

of time to figure out what to eat before they could starve….

And folks like the Brelands found they could not extinguish certain behavior

patterns in the lab no matter what they tried

At first, behaviorists acted like classical physicists at the turn

of the 20th century,

who believed that their current paradigms would

eventually answer things like particles acting like waves…

But eventually, some behaviorists, like the

physicists,

began to realize that new paradigms were needed

But, at the same time that the behaviorists were arguing for

essentially a “tabula rasa” (blank slate),

innate releasing mechanisms and fixed action patterns

ethologists like Lorenz had been focusing on stereotypic behavior

in the wild….

So, there’s the classic story of a stickleback fish….

Males get a red patch when they come into breeding condition, and fight other red-patched males for their territory…

Tinbergen saw a male stickleback go bonkers when it sighted a red

van through a window…

And a male European robin will tear apart a red sock that is

attached to a branch…

Either the animal is really dumb,

or it can’t overcome some innately specified behavior pattern,

or maybe hormones just overcome cognition…

[gray.mov]

So, IRMs and FAPs were initially used to explain issues like

predator avoidance and food preferences and mate choice

But, although some things did seem to be hard-wired…

Additional studies showed that what seemed to be innate mechanisms were often

learned…

albeit very rapidly

So, for example, Hailman showed that the so-called innate

predisposition of newly-hatched gulls to peck at a red dot on their

parents’ beaks

Hatchlings learned within a day that the red spot was a good sign for what to

hit…but they learned

was actually only an innate predisposition to hit a moving

target…

And then there were studies such as Köhler’s on “insight” in

apes…

And although researchers subsequently argued that the animals

may have seen something related that gave them the idea,

and that they definitely had toys that enabled them to gain experience with

stacking….

And that the chimps were actually pretty incompetent…

i.e., they balanced precariously on the corner of the box…

and other researchers had trouble replicating the data…

other studies have indeed shown novel problem-solving abilities

(.e.g., Goodall’s drum-using chimps, crows, parrots)

Weir movie

And if you let a bee explore a field

and then release it from a novel spot in the field

it returns to its hive by the most direct route possible, as tho’ it has figured out a cognitive map of the

field

So, it seemed that neither the innately specified

systems of the ethologists

or the simple rule-driven explanations of the

behaviorists

could fully explain animal behavior

Thus, a number of scientists from the behavioral stance broke rank,

where researchers were beginning to look at issues such as selective

attention, memory, and information processing

and decided to adapt these ideas to the study of animal behavior

and began to look at what had been going on in the realm of human

behavior,

So, with a breakthrough book in the late ’60s, Hulse, Fowler, and

Honig argued that

behavior is best explained by mental representations and information

processing,

and started the so-called “cognitive revolution” in animal behavior

But this was not quite as much of a change as one

might have hoped or expected….

These folks were interpreting their results differently and proposing slightly different

hypotheses,

But they still retained the physical techniques of behaviorism….

That is, locking animals up in what are called ‘operant chambers’ (aka “Skinner boxes”): Studying them

in social isolation

under sterile laboratory conditions and with the same type of tasks (e.g., match-to-sample, oddity, two-choice decisions)

They attempted to, for example, examine whether

animals understood abstract concepts

So they gave the animals tasks like the following where the

correct response lights up and errors darken

*

**

This gives an idea of what the animals had to

uncover…

Choose odd color when objects are backed in white and odd shape if

objects are backed in black…

But is this a concept of same/different?

According to scientists like Premack,

And, moreover, can label exactly what is same or different

an animal that knows ‘same/different’ can transfer to any situation and any modality

and any type of exemplar….

Different color, same shape, same material,

same size

Same color, different shape, different size,

different matter

So, how were scientists really going to understand the abilities

of animals?

Into the fray came Griffin, the man who had determined that bats navigated and hunted via

sonar—

something that was viewed as totally impossible when it was first

proposed

Griffin wrote “The Question of Animal Awareness”, in which he

argued that:

Animals were capable of complex cognitive processing

Such processing likely had to be studied in nature

And that animals were also likely fully conscious

But Griffin did even more than that;

He argued that each anecdote was a data point

And that, if nothing else, should be the basis for a testable hypothesis

he went back a Romanes-like stance, basing many of his arguments on anecdotes

Griffin at the time was ridiculed and dismissed as an aging has-

been

But as we saw from the ant example toward the beginning of

the lecture,

maybe he was on to something…

And, adding even more interest…

or confusion…

to these issues were scientists who were, at this same time in history,

attempting to teach animals to communicate with human

language

Now, remember, Descartes had argued that one reason that animals couldn’t think

was that they couldn’t learn or use human language….

And some researchers had argued…and some still do…that concept formation is impossible

without language

And, of course, such a stance doesn’t even begin to address the issue of whether animals’ natural

communication systems demonstrate possible concept

formation

[and, yes, some evidence suggests they do…e.g.,

neighbor/stranger discriminations]

Given that Griffin argued that animal communication was a

‘window into the mind’….

Was human communication necessary?

Could we develop other ways of questioning animals without using

human language?

Well, the answer is definitely “yes”, although we will see

that even basic forms of ‘language-like’ behavior

simplify the experiments….

As in simply asking what is same or different for two objects…

And, of course, an animal that can ‘talk’ can give

interesting and unexpected answers

Answers that, in fact, suggest a higher level of cognition than

possibly the experimenter expected

But I still haven’t really answered the question of

“What is animal cognition?”

All I’ve done so far is show that over time,

researchers have argued either that animals are dumb

robots

or fully equivalent to humans….

But clearly they are something in between,

and we have to figure out what we are looking for

before we begin our search

There are actually as many definitions of “cognition” as there are researchers in the

field

And we’ll be examining different types of cognition in different

species

We’ll see that different species may be specialized for a particular ability, …

But that are no better than pigeons on match-to-sample tasks in the

lab

like nutcrackers that remember 10,000 caches each

winter

But basically cognition is defined as

an organism’s ability to make a decision by evaluating—or

processing—

current information based on some representation of prior

experience

And, of course, that brings up the definition of a

‘representation’…

but let’s just assume that it exists for now,

and not worry as to whether it is pictorial, verbal, auditory,

etc.

Cognition is thus based on the supposition that the organism does not

react robotically to environmental stimuli,

but rather processes the input and chooses to react in

certain ways

Of course, we have to remember that what seems a logical cognitive process for

humans

And that in itself is another interesting issue

may not be logical for an animal….

If an animal doesn’t much care about the banana slice reward,

but would much rather try to use a tool in a weird interesting way other than the one it has seen work to get the banana slice….

Is it being intelligent or stupid?

Each rake has a hard edge and a floppy rubber edge; the chimp should pull the rake with the food by the hard edge…

Or is the challenge of trying to get the food through a novel action *much* more rewarding than the food

itself?

Thus I’m going to argue that cognitive ability is not simply

the choice of a logical response, but

that the decision-making process include the capacity to choose

from among various possible sets of rules that have been acquired or

taughtthe set that appropriately governs

the current processing of given data

What does this mean and why is it important???

It is important because it means that

organisms limited to examining information

according to a single set of rules (e.g., a matching

procedure)

do not have the chance to exhibit complex cognition

It also means that cognition requires the ability

to transfer a set of skills learned in one context to

another

And, I argue, that many animals in nature need

complex cognitive skills to survive

A foraging bird that has learned to associate green with

unripe/bitter and red with ripe/tasty might do OK in some

instances…

But it also needs to know that for some fruits, red could indicate spoilage and green is optimal

Now, I’ve given just a really brief overview, and there’s a

LOT more to the study of animal cognition…

It is still a relatively new and exciting field, with much to be

learned