The Senses of Insects

Auguste Forel, who in 1909 retired from the Chair of Morbid

Psychology in the University of Zürich, was born on September 1,

1848, and is one of the greatest students of the minds and senses

of the lower animals and mankind. Among his most famous works are

his "Hygiene of Nerves and Mind," his great treatise on the whole

problem of sex in human life, of which a cheap edition entitled

"Sexual Ethics" is published, his work on hypnotism, and his

numerous contributions to the psychology of insects. The chief

studies of this remarkable and illustrious student and thinker for

many decades past have been those of the senses and mental

faculties of insects. He has recorded the fact that his study of

the beehive led him to his present views as to the right

constitution of the state--views which may be described as

socialism with a difference. His work on insects has served the

study of human psychology, and is in itself the most important

contribution to insect psychology ever made by a single student.

Only within the last two years has the work of Forel, long famous

on the European Continent, begun to be known abroad.

_I.--Insect Activity and Instinct_

This subject is one of great interest, as much from the standpoint of

biology as from that of comparative psychology. The very peculiar

mechanism of instincts always has its starting-point in sensations. To

comprehend this mechanism it is essential to understand thoroughly the

organs of sense and their special functions.

It is further necessary to study the co-ordination which exists between

the action of the different senses, and leads to their intimate

connection with the functions of the nerve-centres, that is to say, with

the specially instinctive intelligence of insects. The whole question

is, therefore, a chapter of comparative psychology, a chapter in which

it is necessary to take careful note of every factor, to place oneself,

so to speak, on a level with the mind of an insect, and, above all, to

avoid the anthropomorphic errors with which works upon the subject are

filled.

At the same time the other extreme must equally be

avoided--"anthropophobia," which at all costs desires to see in every

living organism a "machine," forgetting that a "machine" which lives,

that is to say, which grows, takes in nutriment, and strikes a balance

between income and expenditure, which, in a word, continually

reconstructs itself, is not a "machine," but something entirely

different. In other words, it is necessary to steer clear of two

dangers. We must avoid (1) identifying the mind of an insect with our

own, but, above all, (2) imagining that we, with what knowledge we

possess, can reconstruct the mind by our chemical and physical laws.

On the other hand, we have to recognise the fact that this mind, and the

sensory functions which put it on its guard, are derived, just as with

our human selves, from the primitive protoplasmic life. This life, so

far as it is specialised in the nervous system by nerve irritability and

its connections with the muscular system, is manifested under two

aspects. These may be likened to two branches of one trunk.

(_a_) _Automatic_ or _instinctive_ activity. This, though perfected by

repetition, is definitely inherited. It is uncontrollable and constant

in effect, adapted to the circumstances of the special life of the race

in question. It is this curious instinctive adaptation--which is so

intelligent when it carries out its proper task, so stupid and incapable

when diverted to some other purpose--that has deceived so many

scientists and philosophers by its insidious analogy with humanly

constructed machines.

But, automatic as it may appear, instinct is not invariable. In the

first place, it presents a racial evolution which of itself alone

already demonstrates a certain degree of plasticity from generation to

generation. It presents, further, individual variations which are more

distinct as it is less deeply fixed by heredity. Thus the divergent

instincts of two varieties, _e.g._, of insects, present more individual

variability and adaptability than do those instincts common to all

species of a genus. In short, if we carefully study the behaviour of

each individual of a species of insects with a developed brain (as has

been done by P. Huber, Lubbock, Wasmann, and myself, among others, for

bees, wasps, and ants), we are not long in finding noteworthy

differences, especially when we put the instinct under abnormal

conditions. We then force the nervous activity of these insects to

present a second and plastic aspect, which to a large extent has been

hidden from us under their enormously developed instinct.

(_b_) The _plastic_ or _adaptive_ activity is by no means, as has been

so often suggested, a derivative of instinct. It is primitive. It is

even the fundamental condition of the evolution of life. The living

being is distinguished by its power of adaptation; even the amoeba is

plastic. But in order that one individual may adapt itself to a host of

conditions and possibilities, as is the case with the higher mammals and

especially with man, the brain requires an enormous quantity of nerve

elements. But this is not the case with the fixed and specialised

adaptation of instinct.

In secondary automatism, or habit, which we observe in ourselves, it is

easy to study how this activity, derived from plastic activity, and ever

becoming more prompt, complex, and sure (technical habits), necessitates

less and less expenditure of nerve effort. It is very difficult to

understand how inherited instinct, hereditary automatism, could have

originated from the plastic activities of our ancestors. It seems as if

a very slow selection, among individuals best adapted in consequence of

fortunate parentage, might perhaps account for it.

To sum up, every animal possesses two kinds of activity in varying

degrees, sometimes one, sometimes the other predominating. In the lowest

beings they are both rudimentary. In insects, special automatic activity

reaches the summit of development and predominance; in man, on the

contrary, with his great brain development, plastic activity is elevated

to an extraordinary height, above all by language, and before all by

written language, which substitutes graphic fixation for secondary

automatism, and allows the accumulation outside the brain of the

knowledge of past generations, thus serving his plastic activity, at

once the adapter and combiner of what the past has bequeathed to it.

According to the families, _genera_, and species of insects, the

development of different senses varies extremely. We meet with most

striking contrasts, and contrasts which have not been sufficiently

noticed. Certain insects, dragon-flies, for instance, live almost

entirely by means of sight. Others are blind, or almost blind, and

subsist exclusively by smell and taste (insects inhabiting caves, most

working ants). Hearing is well developed in certain forms (crickets,

locusts), but most insects appear not to hear, or to hear with

difficulty. Despite their thick, chitinous skeleton, almost all insects

have extremely sensitive touch, especially in the antennæ, but not

confined thereto.

It is absolutely necessary to bear in mind the mental faculties of

insects in order to judge with a fair degree of accuracy how they use

their senses. We shall return to that point when summing up.

_II.--The Vision of Insects_

In vision we are dealing with a certain definite stimulus--light, with

its two modifications, colour and motion. Insects have two sets of

organs for vision, the faceted eye and the so-called simple eye, or

ocellus. These have been historically derived from one and the same

organ. In order to exercise the function of sight the facets need a

greater pencil of light rays by night than by day. To obtain the same

result we dilate the pupil. But nocturnal insects are dazzled by the

light of day, and diurnal insects cannot see by night, for neither

possess the faculty of accommodation. Insects are specially able to

perceive motion, but there are only very few insects that can see

distinctly.

For example, I watched one day a wasp chasing a fly on the wall of a

veranda, as is the habit of this insect at the end of summer and in the

autumn. She dashed violently in flight at the flies sitting on the wall,

which mostly escaped. She continued her pursuit with remarkable

pertinacity, and succeeded on several occasions in catching a fly, which

she killed, mutilated, and bore away to her nest. Each time she quickly

returned to continue the hunt.

In one spot of the wall was stuck a black nail, which was just the size

of a fly, and I saw the wasp very frequently deceived by this nail, upon

which she sprang, leaving it as soon as she perceived her error on

touching it. Nevertheless, she made the same mistake with the nail

shortly after. I have often made similar observations. We may certainly

conclude that the wasp saw something of the size of a fly, but without

distinguishing the details; therefore she saw it indistinctly. Evidently

a wasp does not only perceive motion; she also distinguishes the size of

objects. When I put dead flies on a table to be carried off by another

wasp, she took them, one after another, as well as spiders and other

insects of but little different size placed by their side. On the other

hand, she took no notice of insects much larger or much smaller put

among the flies.

Most entomologists have observed with what ingenuity and sureness

dragon-flies distinguish, follow, and catch the smallest insects on the

wing. Of all insects, they have the best sight. Their enormous convex

eyes have the greatest number of facets. Their number has been estimated

at 12,000, and even at 17,000. Their aerial chases resemble those of the

swallows. By trying to catch them at the edge of a large pond, one can

easily convince oneself that the dragon-flies amuse themselves by making

sport of the hunter; they will always allow one to approach just near

enough to miss catching them. It can be seen to what degree they are

able to measure the distance and reach of their enemy.

It is an absolute fact that dragon-flies, unless it is cold or in the

evening, always manage to fly at just that distance at which the student

cannot touch them; and they see perfectly well whether one is armed with

a net or has nothing but his hands; one might even say that they measure

the length of the handle of the net, for the possession of a long handle

is no advantage. They fly just out of reach of one's instrument,

whatever trouble one may give oneself by hiding it from them and

suddenly lunging as they fly off. Whoever watches butterflies and flies

will soon see that these insects also can measure the distance of such

objects as are not far from them. The males and females of bees and ants

distinguish one another on the wing. It is rare for an individual to

lose sight of the swarm or to miss what it pursues flying. It has been

proved that the sense of smell has nothing to do with this matter. Thus

insects, though without any power of accommodation for light or

distance, are able to perceive objects at different distances.

It is known that many insects will blindly fly and dash against a lamp

at night, until they burn themselves. It has often been wrongly thought

that they are fascinated. We ought first to remember that natural

lights, concentrated at one point like our artificial lights, are

extremely rare in Nature. The light of day, which is the light of wild

animals, is not concentrated at one point. Insects, when they are in

darkness--underground, beneath bark or leaves--are accustomed to reach

the open air, where the light is everywhere diffused, by directing

themselves towards the luminous point. At night, when they fly towards a

lamp, they are evidently deceived, and their small brains cannot

comprehend the novelty of this light concentrated at one spot.

Consequently, their fruitless efforts are again and again renewed

against the flame, and the poor innocents end by burning themselves.

Several domestic insects, which have become little by little adapted to

artificial light in the course of generations, no longer allow

themselves to be deceived thereby. This is the case with house-flies.

Bees distinguish all colours, and seldom confound any but blue and

green; while wasps scarcely react to differences of colour, but note

better the shape of an object, and note, for instance, where the place

of honey is; so that a change of colour on the disc whereon the honey is

placed hardly upsets them. Further, wasps have a better sense of smell

than bees.

The chief discovery regarding the vision of insects made in the last

thirty years is that of Lubbock, who proved that ants perceive the

ultra-violet rays of the spectrum, which we are unable, or almost

unable, to perceive.

It has lately been proved also that many insects appreciate light by the

skin.

They do not see as clearly as we do; but when they possess

well-developed compound eyes they appreciate size, and more or less

distinctly the contours of objects.

Ants have a great faculty for recognition, which probably testifies to

their vision and visual memory. Lubbock observed ants which actually

recognised each other after more than a year of separation.

_III.--Smell, Taste, Hearing, Pain_

Smell is very important in insects. It is difficult for us to judge of,

since man is of all the vertebrates except the whales, perhaps, the one

in which this sense is most rudimentary. We can evidently, therefore,

form only a feeble idea of the world of knowledge imparted by a smell to

a dog, a mole, a hedgehog, or an insect. The instruments of smell are

the antennæ. A poor ant without antennæ is as lost as a blind man who is

also deaf and dumb. This appears from its complete social inactivity,

its isolation, its incapacity to guide itself and to find its food. It

can, therefore, be boldly supposed that the antennæ and their power of

smell, as much on contact as at a distance, constitute the social sense

of ants, the sense which allows them to recognise one another, to tend

to their larvæ, and mutually help one another, and also the sense which

awakens their greedy appetites, their violent hatred for every being

foreign to the colony, the sense which principally guides them--a little

helped by vision, especially in certain species--in the long and patient

travels which they have to undertake, which makes them find their way

back, find their plant-lice, and all their other means of subsistence.

As the philosopher Herbert Spencer has well pointed out, the visceral

sensations of man, and those internal senses which, like smell, can only

make an impression of one kind as regards space--two simultaneous odours

can only be appreciated by us as a mixture--are precisely those by which

we can gain little or no information relative to space. Our vision, on

the contrary, which localises the rays from various distant points of

space on various distinct points of our retina at the same time, is our

most relational sense, that which gives us the most vast ideas of space.

But the antennæ of insects are an olfactory organ turned inside out,

prominent in space, and, further, very mobile. This allows us to suppose

that the sense of smell may be much more relational than ours, that the

sensations thence derived give them ideas of space and of direction

which may be qualitatively different from ours.

Taste exists in insects, and has been very widely written on, but

somewhat inconclusively. The organs of taste probably are to be found in

the jaws and at the base of the tongue. This sense can be observed in

ants, bees, and wasps; and everyone has seen how caterpillars especially

recognise by taste the plants which suit them.

Much has been written on the hearing of insects; but, in my judgment,

only crickets and several other insects of that class appear to perceive

sounds. Erroneous views have been due to confusing hearing with

mechanical vibrations.

We must not forget that the specialisation of the organ of hearing has

reached in man a delicacy of detail which is evidently not found again

in lower vertebrates.

Pain is much less developed in insects than in warm-blooded vertebrates.

Otherwise, one could not see either an ant, with its abdomen or antennæ

cut off, gorge itself with honey; or a humble-bee, in which the antennæ

and all the front of the head had been removed, go to find and pillage

flowers; or a spider, the foot of which had been broken, feed

immediately on this, its own foot, as I myself have seen; or, finally, a

caterpillar, wounded at the "tail" end, devour itself, beginning behind,

as I have observed more than once.

_IV.--Insect Reason and Passions_

Insects reason, and the most intelligent among them, the social

hymenoptera, especially the wasps and ants, even reason much more than

one is tempted to believe when one observes the regularly recurring

mechanism of their instincts. To observe and understand these

reasonings well, it is necessary to mislead their instinct. Further, one

may remark little bursts of plastic judgment, of combinations--extremely

limited, it is true--which, in forcing them an instant from the beaten

track of their automatism, help them to overcome difficulties, and to

decide between two dangers. From the point of view of instinct and

intelligence, or rather of reason, there are not, therefore, absolute

contrasts between the insect, the mammal, and the man.

Finally, insects have passions which are more or less bound up with

their instincts. And these passions vary enormously, according to the

species. I have noted the following passions or traits of character

among ants: choler, hatred, devotion, activity, perseverance, and

gluttony. I have added thereto the discouragement which is sometimes

shown in a striking manner at the time of a defeat, and which can become

real despair; the fear which is shown among ants when they are alone,

while it disappears when they are numerous. I can add further the

momentary temerity whereby certain ants, knowing the enemy to be

weakened and discouraged, hurl themselves alone in the midst of the

black masses of enemies larger than themselves, hustling them without

taking the least further precaution.

When we study the manners of an insect, it is necessary for us to take

account of its mental faculties as well as of its sense organs.

Intelligent insects make better use of their senses, especially by

combining them in various ways. It is possible to study such insects in

their homes in a more varied and more complete manner, allowing greater

accuracy of observations.

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