Animal behaviour

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ANIMAL BEHAVIOUR

BY
C. LLOYD MORGAN, F.RS.

AUTHOR OF ‘‘ ANIMAL LIFE AND INTELLIGENCE,” ‘‘ HABIT AND INSTINCT,”

“ PSYCHOLOGY FOR TEACHERS,” ETC. ETC.

ILLUSTRATED

LONDON
EDWARD ARNOLD
37 BEDFORD STREET, STRAND
Ig0o

(All rights reserved)

§

ce

PREPACE

My book on “Animal Life and Intelligence” .being out
of print, I undertook to revise it for a new Edition. As
the work of revision proceeded, however, it appeared that
the amended treatment would not fall conveniently under the
previous scheme of arrangement. I therefore decided to -
write a new book under the title of ‘Animal Behaviour.”
A few passages from the older work have been introduced, ~
and some of the observations and conclusions already pub-
lished in greater detail in “ Habit and Instinct” have been
summarized. But it will be found that these occupy a

relatively small space in the following pages.

c. L. M.

University CoLLecr, Brisrox,
October lst, 1900.

CONTENTS

CHAPTER I
ORGANIC BEHAVIOUR

PAGE
Bewavioun In GENERAL . . . . ww. 1
Brnavioun or Conus . ww wwe 3
Corporate BrHaviour. . . . . . . . . . .-. Od
Tue Bewaviour oF Phants . . . . . OD
Rervex Action. . . . . . . ww OB
Tue EvoivTioN or Organic Bewaviour . . . . . . 85

CHAPTER II

CONSCIOUSNESS
Tus Conscious AccoMPANIMENTS oF CERTAIN ORGANIC

CHANGES? 9. G5 gee ae eR See aes ee ge
Tae Harty Sraces or Menran DeveLopment . . . . 48
Later Puases in MentaL DeveLtopmentT. . . . . . 56
Tue EvouuTion or Consciousness . . . . . . . . 421

. CHAPTER III

INSTINCTIVE BEHAVIOUR

DrFiniTion or Instinctive Bewaviour. . . . . . . 63
Instinctive BEHAVIOUR IN INSECTS. . . . . . . . TI
Tue Instinotive Benaviourn or Younec Birps . . . . 84
Tut Conscious Aspect oF INSTINCTIVE Beyaviour . . . 98
Tue Evouution or Instinctive Beuaviour . . . . . 106

vi

CONTENTS

CHAPTER IV
INTELLIGENT BEHAVIOUR

Tue Nature or InreLvicgent BEnAVIOUR .
InvevLicenr Benavrour IN Insects

Some Resutts or ExpErimenT

Tut Evorution or InreL.ticenr Brvaviour
Tue InrLuence or InreLLicence on Insrinct

CHAPTER V
SOCIAL BEITAVIOUR

ImMrra tion ‘

INTERCOMMUNICATION co ntia ja.
SoctaL Communities or Bers and ANTS
ANIMAL TRADITION . a
Tue Evonurion or SoctAL BrHaviour .

CHAPTER VI

THE FEELINGS AND EMOTIONS

Impuusz, Interest, anp Emorion

Puay .

CourtsHip bs OG fas aly Gael Ca, 2
ANIMAL “ AdsruEtics” anp “ Eruics”’ .
Tue Evouution or Freeing snp Exorion

CHAPTER VIL

THE EVOLUTION OF ANIMAL BEHAVIOUR

Tue PuysioLocicaL ASPECT
Tue BrotogicaL Aspecr
THe PsycHoLoGicaL ASPECT
Conrinuity In EvoLurion
InpEx

PAGE

17
123
134
155
168

173
193
205
220
225

235
248
258
270
282

295
305
315
324
338

FIG,

10.

11.

12.

13.

14.

LLEUSTRATIONS

Paramecium. (From “ Animal Biology.” Longmans) .

Behaviour of Paramecia. (After Jennings, American Journal
of Psychology) .

Cell-division. (From “ Animal Biology.” Longmans)

Wapiti with antlers in velvet. (Drawing by Mr. Charles
Whymper, after photograph by Miss Reynolds) .

Wapiti with velvet shredding off. (Drawing by Mr, Charles
Whymper, after photograph by Miss Reynolds) .

Sun-dew leaf and tentacles. (From Darwin's “ Insectivorous
Plants.” Murray. By kind permission of Mr. Francis

Darwin, F-RS,)) «© «© 2°68 8 5 * © © © * & %

Venus’s Fly-trap. (From Darwin’s “Insectivorous Plants.”
Murray. By kind permission of Mr. Francis Darwin, F.R.S.)

Flower of Valisneria

Flower of Catasetum

Flower of Catasetum dissected. (From Darwin’s “ Fertiliza-
tion of Orchids.” Murray. By kind permission of Mr.
Francis Darwin, F.R.S.) . 2... ee eee

Soiitary Wasp stinging Caterpillar. (After Plate III. in Dr.

and Mrs. Peckham’s “Solitary Wasps ”’) :

Solitary Wasp dragging a Caterpillar to its Nest. (After
Plate 1V. in Dr. Peckham’s “Solitary Wasps”) . .

Insect Larve: Sitaris, Argyromeba, and Leucopsis. (After
Fabre “ Souvenirs”) . as Oe a Se

Yucca Flower and Moth

PAGE

17

26

27

28

30

31

75

vill ILLUSTRATIONS

FIG. : PAGE
15. Newly-hatched Chick swimming. (Drawn by Mr. Charles

Whymper, after instantaneous photograpbs and a sketch

by theauthor). 2. 6 6 6 ee ee ee 85
16. Nestling Megapode. (From Dr. R. Bowdler Sharpe’s “ Wonders

of the Bird World.” Wells Gardner). . . . . . . 87

17. Cuckoo ejecting Meadow Pipit. (From Mrs. Hugh Black-
burn’s sketch in “ Birds from Moidart.” David Douglas) . 91

18. Leaf-case of Birch-weevil . . . . . . . . . «. . 121

19. Solitary Wasp using a stone as a tool. (After Plate V. in
Dr. Peckham’s “Solitary Wasps”). . . . . . . « 127

20. Spiders placed by Solitary Wasps in crotches of branching
stems. (After Plate X.in Dr. Peckham’s “Solitary Wasps”) 133

21. Fox-terrier lifting the latch of a gate. (Drawn by Mr. Charles
Whymper, after a photograph by Miss Alice Worsley) . . 145

22. Cage used by Dr. Thorndike. (After figure in “ Animal
Intelligence,” Psychological Review, 1898) . . . . . 148

22, Diagram illustrating Dr. Thorndike’s Experiments. (Based
on data given in his monograph on “ Animal Intelligence”) 150

24. Wood ant. (From Shipley’s “Invertebrates.” A. & C. Black) 207
25. Beetle soliciting food from Ant. (After Wasmann. Enlarged) 213
. 26. Honey-pot Ant. (Enlarged) . 2. © 2. 2... 85

ANIMAL BEHAVIOUR

“ CHAPTER I
ORGANIC BEHAVIOUR

J.— BEHAVIOUR IN GENERAL

We commonly use the word “behaviour” with a wide range
of meaning. We speak of the behaviour of troops in the
field, of the prisoner at the bar, of a dandy in the ball-room.
But the chemist and the physicist often speak of the behaviour
of atoms and molecules, or that of a gas under changing con-
ditions of temperature and pressure. The geologist tells us
that a glacier behaves in many respects like a river, and
discusses how the crust of the earth behaves under the stresses
to which it is subjected. Weather-wise people comment on
the behaviour of the mercury in a barometer as a storm
approaches. Instances of a similar usage need not be multi-
plied. Frequently employed with a moral significance, the
word is at least occasionally used in a wider and more compre-
hensive sense. When Mary, the nurse, returns with the little
Miss Smiths from Master Brown’s birthday party, she is
narrowly questioned as to their behaviour ; but meanwhile
their father, the professor, has been discoursing 'to his students
on the behaviour of iron filings in the magnetic field ; and his
son Jack, of H.M.S. Blunderer, entertains his elder sisters
with a graphic description of the behaviour of a first-class
battle-ship in a heavy sea.
I B

2 ORGANIC BEHAVIOUR

The word will be employed in the following pages in a
wide and comprehensive sense. We shall have to consider, not
only the kind of animal behaviour which implies intelligence,
sometimes of a high order ; not only such behaviour as animal
play and courtship, which suggests emotional attributes ; but
also forms of behaviour which, if not unconscious, seem to lack
conscious guidance and control. We shall deal mainly with
the behaviour of the animal as a whole, but also incidentally
with that of its constituent particles, or cells; and we shall
not hesitate to cite Gn a parenthetic section) some episodes of
plant life as examples of organic behaviour.

Thus broadly used, the term in all cases indicates ait
draws attention to the reaction of that which we speak of as
behaving, in response to certain surrounding conditions or
circumstances which evoke the behaviour. The middy would
not talk of the behaviour of his ship as she lay at anchor in
Portland harbour ; the word is only applicable when there is
action and reaction as the vessel ploughs through a heavy sea,
or when she answers to the helm. Apart from gravitation the
glacier and the river would not “ behave in a similar manner.”
Only under the conditions comprised under the term “ mag-
netic field” do iron filings exhibit certain peculiarities of
behaviour. And so, also, in other cases. The behaviour of cells
is evoked under given organic or external conditions ; instinc-
tive, intelligent, and emotional behaviour are called forth in
response to those circumstances which exercise a constraining
influence at the moment of action.

It is therefore necessary, in a discussion of animal beha-—
viour, that we should endeavour to realize, as far as possible,
in every case, first, the nature of the animal under considera-
tion ; secondly, the conditions under which it is placed ; thirdly,
the manner in which the response is called forth by the cir-
cumstances, and fourthly, how far the behaviour adequately
meets the essential conditions of the situation.

BEHAVIOUR OF CELLS 3

IJ.—BEHAVIOUR OF CELLS

From what has already been said it may be inferred that
our use of the term “ behaviour” neither implies nor excludes
the presence of consciousness. Few are prepared to contend
that the iron filings in a magnetic field consciously group them-
selves in definite and symmetrical patterns, or that sand grains
on a vibrating plate assemble along certain nodal lines because
they are conscious of the effects of the bow by which the
plate is set in sounding vibration. But where organic re-
sponse falls under our observation, no matter how simple and
direct that response may be, there is a natural tendency to
suppose that the behaviour is conscious ; and where the re-
sponse is less simple and more indirect, this tendency is so
strengthened as to give rise to a state of mind bordering on, or
actually reaching, conviction. Nor is this surprising : for, in
the first place, organic responses, even the simplest, are less
obviously and directly related to the interplay of surrounding
circumstances; and, in the second place, they are more
obviously in relation to some purpose in the sense that they
directly or indirectly contribute to the maintenance of life or
the furtherance of well-being. Now where behaviour is com-
plex and subserves an end which we can note and name, there
arises the supposition that it may well be of the same nature
as our own complex and conscious behaviour.

Take for example the behaviour of the Slipper-animalcule,
Paramecium, one of the minute creatures known to zoologists
as Protozoa. The whole animal is constituted by a single cell,
somewhat less than one-hundredth of an inch in length, the
form and behaviour of which may be readily studied under the
microscope. Thousands may be obtained from water in which
some hay has been allowed to rot. The surface of the Para-

-mecium is covered with waving hair-like cilia, by which it is
propelled through the water, while stiffer hairs may be shot out
from the surface at any point where there is a local source of
irritation, as indicated at the top of the accompanying figure.

4 ORGANIC BEHAVIOUR

Two little sacs expand and contract, and serve to drain off
water and waste products from the substance of the cell. Food
is taken in at the end of the funnel, shown in the lower part of
the figure. The cilia here work in such manner as to drive

Fic. 1.—Paramecium.

the particles into and down the tube, and on reaching its inner
end these particles burst through into the semi-fluid sub-
stance, and circulate therein. Just above the funnel there are
two bean-like bodies, the larger of which is known as the
macronucleus, the smaller as the micronucleus.

The process of multiplication is by “fission,” or the
division of each Paramecium into two similar animalcules.
Not infrequently, however, two Paramecia may be seen to
approach each other and come together, funnel to funnel ;
and in each the nuclei undergo curious changes. The macro-
nucleus breaks up, and is scattered. The micronucleus in each
divides into four portions, of which three break up and dis-
appear ; while the fourth again divides into two parts, one to
be retained and the other to be exchanged for the similar
micronuclear product of the other Paramecium. The retained
portion and that received in exchange then unite to form a
new micronucleus. M. Maupas concludes from his careful _
observations that, in the absence of such “ conjugation ” ‘in
the mid=paried of life, Paramecia pass into a state of senility
which ends in decrepitude and death. If this be so, conjuga-
tion is in them necessary for the continuance of a healthy race,

BEHAVIOUR OF CELLS 5

Here we have what a zoologist would describe as a
specialized mode of behaviour of the nuclei; and we have
also the behaviour of the minute creatures (which contain the
nuclei) as they approach each other and come together in
conjugation. Can one wonder that the latter, at any rate, has
been regarded as an example of conscious procedure? In
truth we do not know in what manner and by what subtle
influences the Paramecia are drawn together in conjugation.
But it is scarcely logical to base on such ignorance any posi-
tive assertion as to conscious attraction. It is better to
confess that here is a piece of organic behaviour, the exact
conditions of which are at present unexplained.

We may take from the writings* of Dr. H. 8. Jennings;
of Harvard, some account of other modes of behaviour among
Paramecia. They largely feed upon clotted masses of bacteria.
If a number are placed upon a glass slip, together with a small
bacterial clot, they will be seen to congregate around the clot
and to feed upon it. All apparently press in so as to reach it,
or get as near it as possible. And if a number be placed on
another slide without any clot, they soon collect in groups in
one or more regions, as in Fig. 2,111. It appears as if they were
actuated by some social impulse leading them to crowd together
and shun isolated positions. Nay, more; it seems as if, after
thus collecting and crowding in to some centre of interest, the
attractive influence gradually waned ; the group spreads, and
the Paramecia are less densely packed ; the assembly scatters
more and more, but still seems to be retained by an invisible
boundary beyond which the little creatures do not pass.

Furthermore, if kept in a jar, the Paramecia crowd up
towards the surface where the bacteria clots are floating ; and
if, beneath the cover glass of a slip on which they are under
microscopic examination, a drop of liquid be introduced
through a very fine tube, they will seem either to be attracted
to it, as in Fig. 2, 1., or repelled from it, as in Fig. 2, 11,
according to its nature. From alkaline liquids they are

*. See “The Psychology of a Protozoon,” in the Amer. Jour. of
Psychology, vol. x., No. 4, July, 1899, and the fuller papers there quoted.

6 ORGANIC BEHAVIOUR

repelled ; to slightly acid drops they are attracted, unless the
acidity be too pungent. Heat and cold are alike repellent,
and even a drop of pure distilled water forms an area into
which the Paramecia do not enter.

With such facts before him, the incautious observer may
be led to the conclusion that Paramecia are not only conscious,
but endowed with intelligence and volition. Even M. Binet,*
who occupies a position which should lead him to exercise
more caution, tells us that there is not a single infusorian
which cannot be frightened, and does not manifest its fear by
rapid flight; he speaks of some of these unicellular animals
as “endowed with memory and volition,” and possessed of
“instinct of great precision ;” and he describes the following
stages :—

“(1) The perception of an external object ;

‘“*(2) The choice made between a number of objects ;

(3) The perception of their position in space ;

(4) Movements calculated either to approach the body
and seize it, or to flee from it.” ;

But when we seem to have grasped his point of view,
when we have catalogued the memory, fear, instinct, per-
ception, choice and volition, the whole intelligent edifice
crumbles ; for we are told that ‘we are not in a position to
determine whether these various acts are accompanied by
consciousness, or whether they follow as simple physiological
processes.” ‘To most of us fear, memory, choice, volition,
imply something more than simple physiological processes ;
they imply not only consciousness, but highly elaborated
consciousness.

Dr. Jennings’s researches show that no such implication
can be accepted unless we are prepared to cast aside the
trammels of reasonable caution. In the first place, the whole
matter of feeding appears to be referable to simple organic
behaviour not necessarily involving consciousness. The cilia
in the mouth-groove and funnel constantly wave in such a
manner as to drive a current of water, together with any

* «The Psychic Life of Micro-Organisms,” 1889, p. 61,

BEHAVIOUR OF CELLS 7

particles which float therein, towards the interior ; and the
particles are then engulphed, no matter what their composition
may be. Digestible or indigestible, in they go. There is no
selection of the one or rejection of the other. But, as we
have seen, the Paramecia collect around a bacterial clot and
feed upon it. Surely here there is selection of the nutritious !
Apparently not. They collect in just the same way towards a
piece of blotting-paper, cotton-wool, cloth, sponge, or other
fibrous body, and remain assembled round such an innutritious
centre just as long as round a bacterial clot. There seems to
be no choice in the matter ; contact with any substance gives
rise, a8 an organic response, to the lessening or cessation of
the regular movements in all the cilia except those of the
mouth-groove and funnel. As the Paramecia swim hither and
thither, first one, then another, then more, chance to come in
contact with the bacterial clot, the blotting-paper, or other
substance, and since the lashing of the cilia is then auto-
matically lessened, there they stay ; others find their way to
the same spot in the course of their random movements, and
they, too, stay ; thus many soon collect.

But this does not account for the seemingly social assem-
blages of Paramecia where there is no such substance to arrest
their progress. Dr. Jennings attributes this to the fact that a
dilute solution of carbon-dioxide has, what we may call for the
present, an attractive influence., Ifa bubble of air and a bubble
of carbon dioxide be introduced into the,water in which Para-
mecia are swimming beneath a cover-glass, the animalcules
collect around the carbonic dioxide, but not around the air
bubble. At first they press up close to the bubble of carbon
dioxide, but gradually form a ring farther and farther from
its limiting boundary. This is held to be due to the fact that
it is only the dilute solution of carbonic acid that has the
peculiar “attraction ’’—a stronger solution has a different
effect. And, as the gas dissolves, the Paramecia collect in a
ring just where the solution is sufficiently dilute.

Now carbon dioxide is a product of the organic waste of
living substance ; it is given off by active Paramecia, Where

8 ORGANIC BEHAVIOUR

therefore many are collected together they form a centre of
the production of this substance ; and when other Paramecia
come, in the course of their random movements, into such a

7

Fic. 2.—Behaviour of Paramecia (after Jennings),

centre they remain there and help to swell the numbers in the
cluster. If Paramecia be placed in water to which a dig-
tinctly reddish tinge is given by mixing it with asmall quantity

BEHAVIOUR OF CELLS 9

of rosol—a substance which is decolourized by carbon dioxide,
and is not injurious to Paramecia—it will be seen that, where
the groups are collected, the reddish tinge fades and disappears.
As the groups expand, and are less densely packed, the colour-
less area expands too: and the limits within which the group
is circumscribed are also the limits of decolourization. Dr.
Jennings considers it beyond question that the assembling of
Paramecia is due to the presence in such assemblages of car-
bonic acid produced by the animals themselves. The first
beginning of the crowd may be some small fragment of bacterial
clot or other substance. .

It would seem, then, that Paramecia are attracted by faintly
acid solutions ; and here at least there is, it may be urged, an
element of choice. But even here, according to Dr. Jennings,
there is not only no real choice, but not even any real attrac-
tion. What takes place, according to his observations, is
briefly as follows. Suppose a faintly acid drop be inserted
beneath the cover-glass. Paramecia may almost graze its
boundary without being in any way affected by its presence.
But in their random movements some, and eventually many,
perhaps most, of the little animals chance to enter the faintly
acid region ; but there is no sign of reaction or response ; they
swim on across the drop until they reach its further margin.
Here a reaction does take place. Instead of proceeding on-
wards, slowly revolving on its long axis, a Paramecium thus
situated jerks backwards by a reversal of all the cilia, at the
same time revolving on its axis in a direction opposite to that
in which it was before turning. But the cilia of the mouth-
groove resume their normal mode of working sooner than the
others, and this causes the Paramecium to turn aside. It then
goes ahead until it again reaches the boundary at another point,
when the same behaviour is seen. The course of such a
Paramecium is shown in Fig. 2, Iv.

If, instead of a faintly acid drop, a little alkaline liquid be
introduced beneath the cover-glass, the Paramecium similarly
jerks backward and turns aside on reaching its outer boundary.
The turning may carry it away from the alkali, as shown in

10 ORGANIC BEHAVIOUR

Fig. 2, v.; but it just as often brings it again towards the
drop, especially a large one. It seems to be a matter of
chance which result follows. But eventually the little crea-
ture sails off, since each time it comes within the influence
of the alkaline fluid it jerks back and turns. It appears,
then, that when it is swimming in a normal solution a faintly
acid liquid does not much modify its behaviour, but an alka-
line fluid evokes a reversal of the cilia; and that when it
isa slightly acid solution, not only does stronger acid cause
reversal, but normal fluid producesa similar result. A reaction
of essentially the same kind is in fact called forth by such
different stimuli as chemical substances, water heated above
the normal temperature, or cooled considerably below it, and
fluids which cause changes of internal pressure within the
substance of the cell, Nor does it matter where the stimulus
is applied. If it be applied at the hinder end the infusorian
still jerks backward, though this may drive it into a destructive
solution and thus cause death. There is, however, some evi-
dence of different behaviour in some infusorians according as
the stimulus is here or there. In other words, the behaviour
is to some extent related to the position of the part stimulated.

Furthermore, it may be gathered from Dr. Jennings’s ac-
count that there is nothing to lead us to suppose that such free
living cells show any indication of what may be regarded as
the keynote of intelligent behaviour. They do not profit by
experience. ‘T'hey exhibit organic reactions which may be ac-
companied by some dim form of consciousness, but which do not
seem to be under the guidance of such consciousness, if it exist.

One of the first lessons which the study of animal behaviour,
in its organic aspect, should impress upon our minds is, that
living cells may react to stimuli in a manner which we per-
ceive to be subservient to a biological end, and yet react
without conscious purpose—that is to say, automatically. The
living cell assimilates food and absorbs oxygen, it grows and
subdivides, it elaborates secretions, produces a skeletal frame-
work or covering, rids itself of waste products, responds to stimuli
in a definite fashion, moves hither and thither at random, its

BEHAVIOUR OF CELLS TL

functional activities being stimulated or checked by many
- influences ; and yet this varied life may give no evidence of a
guiding consciousness : if purpose there be, it lies deeper than
its protoplasm, deeper than the dim sentience which may be
present or may be absent—we cannot tell which.

And when the cells are incorporated in the body of one of
the higher animals, instead of each preserving a free and
nomad existence; when they become the multitudinous con-
stituents of an organic republic with unity of plan and unity
of biological end, then the behaviour of each is limited in range
but perfected within that range, in subservience to the require-
ments of the more complex unity. The muscle cell contracts,
the gland-cell secretes, the rods and cones of the retina respond
to the waves of light, and all the normal responses of the
special cells go on with such orderly regularity that the term
behaviour seems scarcely applicable to reactions so stereotyped.
But the physiologist and the physician know well that such
uniformity of response is dependent on uniformity of conditions.
A little dose of some drug will profoundly modify and render
abnormal the procedure which was before so mechanical in its
exactitude ; and we are thus led to see how dependent the
orderly behaviour really is on the maintenance of certain sur-
rounding conditions.

Moreover, the existence of every cell in the body corporate
is the outcome of a process of division involving a special mode
of behaviour in the nucleus, of which we are only beginning to
euess the meaning and significance, and of which we seek in
vain to find an explanation in mechanical terms. And when
we trace these divisions back to their primary source in the
fertilized ovum, we find changes and evolutions in the nuclear
matter of which it can only be said that the more they are
studied the more complex and varied do they appear.

The egg, or ovum, is a single cell produced by the female,
and varying much in size, according to the amount of
food-yolk with which it is supplied. Like other cells, it has
a nucleus, and this undergoes changes which are definitely
related to the fertilization of the ovum, which we describe as

12 ORGANIC BEHAVIOUR

the biological end. Such preparatory changes for a future
contingency are especially characteristic of organic behaviour.
There is nothing like it in the mineral kingdom. The nucleus
divides into two parts, one of which passes out of the ovum
and is lost. The nucleus again divides, and again one part passes
out and is lost. Thus only one quarter of the original amount
of nuclear matter remains. Now, division of the nucleus occurs
whenever an animal cell divides ; but in this case (apart from
details which would here be out of place) there is this difference.
During the ordinary division of cells there are found in the
nucleus a definite number of curved rods, and this number is
constant for any given species; but in the nucleus which re-
mains in the ovum after three parts of its substance are lost,
the number of rods has been reduced to half that which is
common to the species. The egg is now ready for fertilization.
A minute active cell, which is produced by the male, and
which also has only half the normal number of rods, enters
the ovum. The two nuclei approach each other, and give rise
to the single nucleus of the fertilized ovum, which thus has
the full number of rods—half of them derived from one
parent, half from the other parent. The sperm cell of the
male adds little to the store of protoplasm in the ovum ; but
it introduces a minute body, which seems to initiate sub-
sequent divisions of the cell. The nature of these divisions
may be seen in the accompanying diagrammatic figure. Ina
the cell is just preparing to divide. Above the nucleus is the
minute body (centrosome) just spoken of, which has already
divided. In the nucleus the matter of which the rods will
be constituted is net-like. In 8B this net-work has taken on
the new form of a coiled thread, while the divided body above
is associated with a spindle of delicate fibres. In c the
membrane round the nucleus has disappeared, and the coiled
thread has broken up into curved rods (chromosomes), four
of which are shown. The two halves of the minute body
_ form the centres of radiating stars. In p each curved rod
has split along its length, and the two parts are being drawn
asunder towards the centres of the two stars; the cell itself is

BEHAVIOUR OF CELLS 13

beginning to divide. In = the process is carried a step
further, while in F the cell has completely divided into two :
the rods have disappeared as such, and are replaced by a net-
work; a new nuclear membrane has been formed, and the
- minute body has again divided preparatory to the further
division of the cell.

E~ A
Fie. 3.—Cell-division.

Such, stripped as far as possible of technicalities, are some
of the facts concerning the behaviour of cells and their nuclei
during the process of cell-multiplication. No good purpose
would be subserved by pretending that we fully understand
them. The splitting of the rods does indeed seem an efficient
means to the end of securing a fair division of the nuclear
substance, which, according to many biologists, is the organic
bearer of hereditary qualities in the cells. But that is nearly
all that we can say. Is the process accompanied by some form
of sentience? We do not know. That it is controlled and
guided by any consciousness in the cell is most improbable. But
if it be a purely organic and unconscious process it should at
least impress on our minds the fact that such organic be-
haviour may reach a high degree of delicacy and complexity.

14 ORGANIC BEHAVIOUR

III. Corporate BreHAvIOUR

The word “corporate” is here applied to the organic
behaviour of cells when they are not independent and free,
but are incorporated in the animal body, and act in relation to . .
each other. If the behaviour of the individual cell during
division impresses us with the subtle intricacy of organic
processes, the behaviour of the growing cell-republic during
the early stages of organic development must impress us’ no
less forcibly. We place the fertilized egg of a hen in an incu-
bator, and supply the requisite conditions of warmth, moisture,
and fresh air. Before the egg is laid cell-division has begun.
A small patch of closely similar cells has formed on the surface
of the yolk. Further subdivision is then arrested until the
warmth of incubation quickens again the patch into life. But
when once thus quickened no subsequent temporary arrest is
possible—life will not again lie dormant. If arrest there be
it is that of death. And from that little patch of cells, which
spreads further and further over the yolk, a chick is developed.
Into the intricate technicalities of embryology this is not
the place to enter. But it isa matter of common knowledge
that, whereas we have to-day an egg such as we eat for break-
fast, three weeks hence we shall have a bright active bird, a
cunningly wrought piece of mechanism, and, more than that, a
going machine. During this wonderful process the cellular
constituents take on new forms and perform new functions,
all in relationship to each other, all as part of one organic
whole. Here bones are developed to form a skeletal frame-
work, there muscles are constituted which shall render orderly
movements possible ; feathers, beak, and claws take shape as
products of the skin ; gut and glands prepare for future modes
of nutrition ; heart and blood-vessels undergo many changes,
some reminiscent of bygone and ancestral gill-respiration,
some in relation to the provisional respiration of the embryo
by means of a temporary organ that spreads out beneath the
shell, some preparatory to the future use of the lungs,—some,
again, related to the absorption of food from the yolk, others

CORPORATE BEHAVIOUR 15

to subsequent means of digestion; nerve, brain, and sense-
organs differentiate. A going machine in the egg, the chick .
is hatched, and forthwith enters on a wider field of behaviour.
Few would think of attributing to the consciousness of the
embryo chick any guiding influence on the development of its
bodily structure, any control over the subtle changes and dis-
positions of its constituent cells. But no sooner does the
chick, when it is hatched, begin to show wider modes of in-
stinctive behaviour, than we invoke conscious intelligence for
their explanation, seemingly forgetful of the fact that there is
no logical ground for affirming that, while the marvellous
delicacies of structure are of unconscious organic origin, the
early modes of instinctive behaviour are due to the guidance
of consciousness. Such modes of behaviour will, however, be
considered in another chapter. Here we have to notice that
the unquestionably organic behaviour of the incorporated re-
public of cells may attain to a high degree of complexity, and
may serve a distinctly biological end.

There is, perhaps, no more striking instance of rapid and
vigorous growth than is afforded by the antlers of deer,* which
are shed and renewed every year. In the early summer, when
growing, they are covered over with a dark hairy skin, and are
said to be “‘in velvet.” If you lay your hand on the growing
antler, you will feel that it is hot with the nutrient blood that
is coursing beneath it. It is, too, exceedingly sensitive and
tender. An army of tens of thousands of busy living cells is
at work beneath that velvet surface, building the bony antlers,
preparing for the battles of autumn. Each minute cell, working
for the general good, takes up from the nutrient blood the
special materials it requires ; elaborates the crude bone-stuff,
at first soft as wax, but ere long to become hard as stone ; and
then, having done its work, having added its special morsel to
the fabric of the antler, remains embedded and immured, buried
beneath the bone-products of its successors or descendants.
No hive of bees is busier or more replete with active life than

* This paragraph is taken from “Animal Life and Intelligence,”
p. 28.

16 ORGANIC BEHAVIOUR

the antler of a stag as it grows beneath the soft, warm velvet.
And thus are built up in the course of a few weeks those
splendid “ beams,” with their “tynes” and “snags,” which, in
the case of the wapiti, even in the confinement of our Zoological

Fic. 4.--Wapiti with antlers in velvet.

Gardens, may reach a weight of thirty-two pounds, and which,
in the freedom of the Rocky Mountains, may reach such a size
that a man may walk, without stooping, beneath the archway
made by setting up upon their points the shed antlers. When
the antler has reached its full size, a circular ridge makes its

CORPORATE BEHAVIOUR 17

appearance at a short distance from the base. This is the
“burr,” which divides the antler into a short “pedicel” next
the skull, and the “beam” with its branches above. The
circulation in the blood-vessels of the beam now begins to
languish, and the velvet dies and peels off, leaving the hard,

Fic. 5.—Wapiti with velvet shredding off.

bony substance exposed. Then is the time for fighting, when

the stags challenge each other to single combat, while the

hinds stand timidly by. But when the period of battle is over,

and the wars and loves of the year are past, the bone beneath

the burr begins to be eaten away, through the activity of

certain large bone-absorbing cells, and, the base of attachment
C

18 ORGANIC BEHAVIOUR

being thus weakened, the antlers are shed ; the scarred surface
skins over and heals, and only the hair-covered pedicel of the
antler is left.

We have no reason to suppose that this corporate cellular

behaviour, involving the nicely adjusted co-operation of so vast
an army of organic units, is under the conscious guidance of
the stag. And yet how orderly the procedure ! how admirable
_ the result! Nor is there an organ or structural part. of the
stag or any other animal that does not tell the same tale.
This is but one paragraph of the volume in which is inscribed
the varied and wonderful history of organic behaviour in its
corporate aspect. Is it a matter for wonder that the cause of
such phenomena has been regarded as “a mystery transcending
naturalistic conception ; as an alien influx into nature, baffling
scientific interpretation” ? And yet, though not surprising,
this attitude of mind, in face of organic phenomena, is illogical,
and is due partly to a misconception of the function of scientific
interpretation, partly to influences arising from’ the course
pursued by the historical development of scientific knowledge.
The function of biological science is to formulate and to
express in generalized terms the related antecedences and
sequences which are observed to occur in animals and plants.
This can already be done with some approach to precision.
But the underlying cause of the observed phenomena does not
fall within the purview of natural science ; it involves meta-
physical conceptions. It is no more (and no less) a “ mystery ”
“than all causation in its last resort—as the raison d’étre of
observed phenomena—is a mystery. Gravitation, chemical
affinity, crystalline force,—these are all ‘‘ mysteries.”

If the mystery of life, lying beneath and behind organic
behaviour, be said to baffle scientific interpretation, this is
because it suggests ultimate problems with which science as
such should not attempt to deal. The final causes of vital
phenomena (as of other phenomena) lie deeper than the probe ©
of science can reach. But why is this sense of mystery
especially evoked in some minds by the contemplation of
organic behaviour, by the study of life? Partly, no doubt,

CORPORATE BEHAVIOUR 19

because the scientific interpretation of organic processes is but
recent, and in many respects incomplete. People have grown
so accustomed to the metaphysical assumptions employed by
physicists and chemists when they speak of the play of
_ crystalline forces and the selective affinities of atoms, they
have been wont for so long to accept the “mysteries” of
crystallization and of chemical union, that these assumptions
have coalesced with the descriptions and explanations of
science ; and the joint products are now, through custom,
cheerfully accepted as natural. Where the phenomena of
organic behaviour are in question, this coalescence has not yet
taken place ; the metaphysical element is on the one hand
proclaimed as inexplicable by natural science, and on the other
hand denied even by those who talk glibly of physical forces as
the final cause of the phenomena of the inorganic world.

So much reference to the problems which underlie the
problems of science seems necessary. It is here assumed that
the phenomena of organic behaviour are susceptible of scientific
discussion and elucidation. But even assuming that an
adequate explanation in terms of antecedence and sequence
shall be thus attained by the science of the future, this will
not then satisfy, any more than our inadequate explanations
now satisfy, those who seek to know the ultimate meaning and
reason of it all: What makes organic matter behave as we. see
it behave ? what drives the wheels of life, as it drives the
planets in their courses ? what impels the egg to go through its
series of developmental changes? what guides the cells along
the divergent course of their life-history ? These are questions
the ultimate answers to which lie beyond the sphere of science
—questions which man (who is a metaphysical being) always
does and always will ask, even if he rests content with the
answer of agnosticism ; but questions to which natural science
never will be able, and should never so much as attempt, to
give an answer.

Enough has now been said to show that organic behaviour
is a thing swi generis, carrying its own peculiar marks of dis-
tinction ; and further, that, for science, this is just part of the

20 ORGANIC BEHAVIOUR

constitution of nature, neither more nor less mysterious than,
let us say, crystallization or chemical combination. But
associated and closely interwoven with all that is distinctively
organic there is much which can to some extent be interpreted
in terms of physics and chemistry.

The animal * has sometimes been likened to a steam-engine,
in which the food is the fuel which enters into combustion
with the oxygen taken in through the lungs. It may be worth
while to modify and modernize this analogy—always remember-
ing, however, that such an analogy must not be pushed too far.

In the ordinary steam-engine the fuel is placed in the fire-
box, to which the oxygen of the air gains access; the heat
produced by the combustion converts the water in the boiler
into steam, which is made to act upon the piston, and thus set
the machinery in motion. But there is another kind of engine,
now extensively used, which works on a different principle.
In the gas-engine the fuel is gaseous, and it can thus be
introduced in a state of intimate mixture with the oxygen with
which it is to unite in combustion. This isa great advantage.
The two can unite rapidly and explosively. In gunpowder the
same end is effected by mixing the carbon and sulphur with
nitre, which contains the oxygen necessary for their explosive
combustion. And this is carried still further in dynamite and
gun-cotton, where the elements necessary for explosive com-
bustion are not merely mechanically mixed, but are chemically
combined in a highly unstable compound.

But in the gas-engine, not only are the fuel and the oxygen
thus intimately mixed, but the controlled explosions are caused
to act directly on the piston, and not through the intervention
of water in a boiler. Whereas, therefore, in the steam-engine
the combustion is to some extent external to the working of
the machine, in the gas-engine it is to a large extent internal
and direct.

Now, instead of likening the animal as a whole to a steam-
engine, it is more satisfactory to liken each cell to an automatic

* The following paragraphs are taken with some slight changes from
“ Animal Life and Intelligence,” pp. 30-35.

CORPORATE BEHAVIOUR 21

gas-engine which manufactures its own explosive. During
the period of repose which intervenes between periods of
activity, its protoplasm is busy in construction, taking from
the blood-discs oxygen, and from the blood-fluid carbonaceous
and nitrogenous materials, and knitting these together into
relatively unstable explosive compounds, which play the part
of the mixed air and gas of the gas-engine. A resting muscle
may be likened to a complex and well-organized battery of
gas-engines. On the stimulus supplied through a nerve-
channel a series of co-ordinated explosions takes place: the
gas-engines are set to work; the muscular fibres contract ;
the products of the silent explosions are taken up and hurried
away by the blood-stream; and the protoplasm prepares a
fresh supply of explosive material. Long before the invention
of the gas-engine, long before gun-cotton or dynamite were
dreamt of, long before some Chinese or other inventor first
mixed the ingredients of gunpowder, organic nature had utilized
the principle of controlled explosions in the protoplasmic cell,
and thus rendered animal behaviour possible.

Certain cells are, however, more delicately explosive than
others. Those, for example, on or near the external surface
of the body—those, that is to say, which constitute the end-
organs of the special senses—contain explosive material which
may be fired by a touch, a sound, an odour, the contact with
a sapid fluid or a ray of light. The effects of the explosions
in these delicate cells, reinforced in certain neighbouring nerve-
batteries, are transmitted down the nerves as waves of subtle
chemical or electrolytic change, and thus reach that wonderful
aggregation of organized and co-ordinated explosive cells, the
brain. Here it is again reinforced and directed (who, at
present, can say how ?) along fresh nerve-channels to muscles,
or glands, or other organized groups of explosives. And in
the brain, somehow associated with the explosion of its cells,
consciousness, the mind-element, emerges ; of which we need
only notice here that it belongs to a wholly different order of
being from the physical activities and products with which we
are at present concerned.

22 ORGANIC BEHAVIOUR

We must not press the explosion analogy too far. The
essential thing seems to be that the protoplasm of the cell has
the power of building up complex and unstable chemical
compounds, which are perhaps stored in its spongy sub-
stance; and that these unstable compounds, under the
influence of a stimulus (or, possibly, sometimes spontaneously),
break down into simpler and more stable compounds. In the
case of muscle-cells, this latter change is accompanied by an
alteration in length of the fibres, aud consequent movements
in the animal, the products of the disruptive change being
useless or harmful, and being, therefore, removed as soon as
possible. But very frequently the products of explosive activity
are made use of. In the case of bone-cells, one of the products
of disruption is of permanent use to the organism, and con-
stitutes the solid framework of the skeleton. In the case of
the secreting cells in the salivary and other digestive glands,
some of the disruptive products are of temporary Value for
the preparation of the food. It is probable that these useful
products of disruption, permanent or temporary, took their
origin in waste products for which natural selection has found
a use, and which have been gradually rendered more and more
efficacious in modes of organic behaviour increasingly complex.

In the busy hive of cells which constitutes what we call
the animal body, there is thus ceaseless activity. During
periods of apparent rest the protoplasm is engaged in con-
structive work, building up fresh supplies of unstable matertals,
which, during periods of apparent activity, break up into
simpler and more stable substances, some of which are useful
to the organism, while others must be got rid of as soon
as possible. From another point of view, the cells during
apparent rest are storing up energy to be utilized by the
animal during its periods of activity. The storing up of
available energy may be likened to the winding up of a watch
or clock; it is when an organ is at rest that the cells are
winding themselves up; and thus we have the apparent
paradox that the cell is most active and doing most work
when the organ of which it forms a part is at rest. During

CORPORATE BEHAVIOUR 23

the repose of an organ, in fact, the cells are busily working
in preparation for the manifestation of energetic action that
is to follow. Just as the brilliant display of intellectual
activity in a great orator is the result of the silent work of
a lifetime, so is the physical manifestation of muscular power
the result of the silent preparatory work of the muscle-cells.

It may, perhaps, seem strange that the products of cellular
life should be reached by the roundabout process of first
producing unstable compounds, from which are then formed
more stable substances, useful for permanent purposes as in
bone, or temporary purposes as in the digestive fluids. It
seems a waste of power to build up substances unnecessarily
complex and stored with an unnecessarily abundant supply of
energy. But only thus could the organs be enabled to act
under the influence of stimuli, and afford examples of corporate
behaviour. They are like charged batteries ready to discharge
under the influence of the slightest organic touch. In this
way, too, is afforded a means by which the organ is not
dependent only upon the products of the immediate activity
of the protoplasm at the time of action, but can utilize the
store laid up during preceding periods of rest.

Sufficient has now been said to illustrate the nature of
some of the physical processes which accompany organic
behaviour in its corporate aspect. The fact that should
stand out clearly is that the animal body is stored with large
quantities of available energy resident in highly complex and
unstable chemical compounds, elaborated by the constituent
cells. These unstable compounds, eminently explosive accord-
ing to our analogy, are built up of materials derived from
two different sources—from the nutritive matter (containing
carbon, hydrogen, and nitrogen) absorbed during digestion,
and from oxygen taken up from the air during respiration.
The cells thus become charged with energy that can be set
free on the application of the appropriate stimulus, which may
be likened to the spark that fires the explosive.

Let us note, in conclusion, that it is through the blood-
system, ramifying to all parts of the body, and the nerve-

24 ORGANIC BEHAVIOUR

system, the ramifications of which are not less perfect, that
one of the larger and higher animals is knit together into
an organic whole. The former carries to the cell the raw
materials for the elaboration of its explosive products, and,
after the explosions, carries off the waste products which result
therefrom. The nerve-fibres carry the stimuli by which the
explosive is fired, while the central nervous system organizes,
co-ordinates, and controls the explosions, and initiates the
elaboration of the explosive compounds. Blood and nerves
co-operate to render corporate behaviour possible.

IV.—THE BEHAVIOUR OF PLANTS

A short parenthetic section on the behaviour of plants may
serve further to illustrate the nature of organic behaviour.
We have seen that Paramecium is apparently attracted by
faintly acid solutions, and have briefly considered Dr. Jennings’s
interpretation of the facts disclosed by careful observation.
In the ferns the female element, or ovum, is contained in a
minute flask-shaped structure (archegonium), in the neck and
mouth of which mucilaginous matter, with a slightly acid
reaction, is developed ; and this is said to exercise an attractive
influence on the freely swimming ciliated male elements, or
spermatozoids, which are necessary for fertilization. ‘* Now,
it has been shown by experiment that the spermatozoids of
ferns are attracted by certain chemical substances, and especially
by malic acid. If artificial archegonia are prepared (consisting
of tiny capillary glass-tubes) and filled with mucilage to which
a small quantity of this acid has been added, they are found,
when placed in water containing fern-spermatozoids, to exercise
the same attraction upon them which the real archegonia
exercise in nature. The malic acid gradually diffuses out into
the water, and the spermatozoids are influenced by it, so that
they move in the direction in which the substance is more
concentrated, 7.e, towards the tube. Although it cannot be
proved that the archegonia themselves contain malic acid, as
they are too small for a recognizable quantity to be obtained

THE BEHAVIOUR OF PLANTS 25

from them, yet there can be little doubt that the natural
archegonia owe their attractive influence to the same chemical
agent which has proved efficacious in experiment.”* In the
light of Dr. Jennings’s observations, it is perhaps not im-
probable that this so-called attractive influence is similar to
that seen in Paramecium ; and that the spermatozoids enter
the organic acid in the course of their random movements, and
there remain. Be that as it may, the male elements collect in
the mucilaginous mass, and pass down the neck of the flask
until one reaches and coalesces with the female element, or
ovum, and effects its fertilization. Here we have organic
behaviour unmistakably directed to a biological end—behaviour
which may indeed be accompanied by some dim form of
consciousness, but which is due to a purely organic reaction.
It is scarcely satisfactory to say that the spermatozoids “ possess
a certain power of perception, by which their movements are
guided.” + If consciousness be present, it is probably merely
an accompaniment of the response, and has no directive in-
fluence on its nature and character.

In the higher plants, as in the higher animals, the
differentiation and the orderly marshalling of the cell-progeny
arising from the coalescent male and female elements, afford,
during development, examples of corporate organic behaviour
which can be more readily described than explained, but which
not less clearly subserve definite biological ends, and in many
cases, such as the direction of growth in radicles and roots, the
curling of tendrils, and the reaction to the influence of light
and warmth, are related to and evoked by the environing con-
ditions. More closely resembling familiar modes of behaviour
in animals are such movements as are seen in the “ tentacles”
which project from the upper surface and margin of the Sun-
dew leaf. Their knobbed ends secrete a sticky matter, which
glistens in the sun, and to which small foreign bodies readily
adhere. If particles of limestone, sand, or clay, such as may

* D. H. Scott, “An Introduction to Structural Botany,” part ii.,
“ Flowerless Plants,” pp. 70, 71.
+ Ibid., p. 71.

26 ORGANIC BEHAVIOUR

be blown by the wind, touch and stick to these knobs, there
follows an exudation of acid liquid, but no marked and con-
tinuous change occurs in the position of the tentacles. But
should an insect alight on the leaf, or a small piece of meat be
placed upon the tentacles, not only is there a discharge of acid
juice, but a ferment is also produced, which has a digestive
action on the nitrogenous matter.
Slowly the tentacle curves inwards
and downwards, as one’s finger
may bend towards the palm of
one’s hand; neighbouring ten-
tacles also turn towards and
incline on to the stimulating
substance ; then others, further
off, behave in a similar way, until
all the tentacles, some’ two hun-
dred in number, are inflected and
converge upon the nitrogenous
particle. Nay, more: ‘“ When
two little bits of meat are placed
simultaneously on the right and
re eee me es left halves of the same Sun-dew
Leaf (enlarged) with the ten. leaf, the two hundred tentacles
tacles on one side inflected divide into two groups, and each
over a bit of meat placed on : : :
the disc. (From Darwin’s O2€ of the groups directs its aim
“ Insectivorous Plants.”) to one of the bits of meat.” *
The movements, though slow, are orderly, methodical, and
effective, the secretions of many glands being brought to bear
on just those substances which are capable of digestion and
absorption by the plant. The seemingly concerted action is
moreover due to an organic transmission of impulses from cell
to cell—a transmission accompanied by visible changes in a
purple substance contained within the cells. In the Sun-dew
any tentacle may form the starting-point of the spreading
wave of impulse. But in the Venus’s Fly-trap there are six

* Kerner, “Natural History of Plants,” translated by F. W. Oliver
>
vol. i, p. 145.

THE BEHAVIOUR OF PLANTS 27

delicate spines, the slightest touch on any one of which causes
the two halves of the specially modified leaf-end to fold inwards
on the midrib as a hinge. ‘The transmission of impulse is
more rapid, the trap closing in a few seconds ; and electric
currents have been observed to accompany the change. Tooth-
like spines at the edge of the trap interlock, and serve to
prevent the escape of small insects, while short-stalked purple
glands secrete an acid digestive juice. Division of labour has
been carried further; and organic behaviour, not less pur-
posive, is carried out in a manner even more effective.

Fic. 7.—Venus’s Fly-trap (Dionza). Leaf viewed laterally in its ex-
panded state. (From Darwin’s “ Insectivorous Plants.”)

In other plants adaptive movements are well known.
“Few phenomena have such a peculiar appearance as the
movements which occur in the sensitive Oxalis when rain comes
on. Not only do the leaflets on which the finest rain-drops
fall fold together in a downward direction, but all the neigh-
bouring ones perform the same movement, although they have
not themselves been shaken by the impact of the falling drops.
The movement is continued to the common leaf-stalk bearing
the numerous leaflets. This also bends down towards the
ground. The rain-drops now slide over the bent leaf-stalk and
down over the depressed leaflets, and not a drop remains
behind on their delicate surfaces.” * The waves of impulse
are said to be transmitted along definite lines, and to cause

* Kerner, “ Natural History of Plants,” vol. i., p. 536.

28 ORGANIC BEHAVIOUR

the expulsion of water from certain cells at the point of
insertion of the leaflets or leaf-stalks, rendering them flaccid.
Appealing even more strongly to the popular imagination,
though probably not of deeper biological significance, is the
behaviour of plants in relation to the essential process of
fertilization. Only two examples can here be cited. Valisneria
spiralis is an aquatic plant, with long submerged strap-like
leaves, which grows in still water in Southern Europe. The
female flower is enclosed in two translucent bracts, which
form a protective bladder so long as the flower is beneath

Fic. 8.—F lower of Valisneria.

the surface of the water; but the flower-stalk continues to
grow until the flower reaches the surface, when it becomes
freely exposed by the splitting of the bracts. There are three
boat-shaped sepals, which act as floats; three quite minute
petals ; and three large fringed stigmas, which project over the
abortive petals in the space between the boat-like sepals. The
flower is now ready for fertilization.

The male flowers, which are developed on different indi-
viduals from those which produce the female flowers, grow in
bunches beneath an investing bladder. The stalk does not
clongate, so that the bladder never rises far above the bottom,
and remains completely submerged. Here the bladder bursts,
and the male flowers, with short stalks, are detached. Each
has three sepals, which enclose and protect the stamens. The
separated flower now ascends to the surface, the sepals open

THE BEHAVIOUR OF PLANTS 29

and form three hollow boats, by means of which the flower
floats freely, while the two functional stamens project upwards
and somewhat obliquely into the air, exposing the large sticky
pollen-cells. Blown hither and thither by the wind, these
little flower-boats “accumulate in the neighbourhood of fixed
bodies, especially in their recesses, where they rest like ships
in harbour. When the little craft happen to get stranded
in the recesses of a female Valisneria flower, they adhere to
the tri-lobed stigma, and some of the pollen-cells are sure to
be left sticking to the fringes on the margins of the stigmatic
surface.” * ;

This is a good example of purely organic behaviour ad-
mirably adapted to secure a definite and important biological
end. Few will be likely to contend that it is even accom-
panied by, still less under the guidance of, any conscious fore-
sight on the part of the plant. And the lesson it should teach
is that, in the study of organic behaviour, adaptation to the
conditions of existence is not necessarily the outcome of
conscious guidance.

It is well known that the orchids exhibit, in their mode
of fertilization, remarkable adaptations by which the visits of
insects are rendered. subservient to the needs of the plant. In
the Catasetums, for example, the male flower may be described
as consisting of two parts—a lower part, the cup-like labellum
(Figs. 9, 1), which constitutes a landing-stage on which insects
may alight; and an upper part, the column (Fig. 9, ¢),
surrounded by the upper sepal and petals. In the upper part
of the column the pollen-masses are borne at one end of an
elastic pedicel, at the other end of which is an adhesive disc,
and the rod is bent over a pad so as to be in a state of strain.
The disc is retained in position by a membrane with which
two long tubular horns (Figs. 9,4; 10, a) are continuous.
These project over the labellum, where insects alight to
gnaw its sweet fleshy walls, and if they be touched, even very
lightly, they convey some stimulus to the membrane which
surrounds and connects the disc with the adjoining surface,

* Kerner, “Natural History of Plants,” vol. ii., p. 182.

3° ORGANIC BEHAVIOUR

causing it instantly to rupture ; and as soon as this happens,
the disc is suddenly set free. The highly elastic pedicel
then flirts the disc out of its chamber with such force that the
whole is ejected, sometimes to a distance of two or three feet,

Fic. 9.—Flower of Catasetum; c, column; h, horns; 7, labellum.

bringing away with it the two pollen-masses. ‘The utility
of so forcible an ejection is to drive the soft and viscid cushion
of the dise against the hairy thorax of the large hymenopterous
insects which frequent the flowers. When once attached to
an insect, assuredly no force which the insect could exert
would remove the disc and pedicel, but the caudicles [by which
the pollen-masses are attached] are ruptured without much
difficulty, and thus the balls of pollen might readily be left on
the adhesive stigma of the female flower.” *

Here again we have adaptive behaviour of exquisite nicety,
and we have the transmission of an impulse very rapidly along
the cells of the irritable horns, followed by the sudden rupture
of a membrane. Beautiful, however, as is the adaptation,
effective as it is to a definite biological end, the organic
behaviour does not afford any indication of the guidance of

* Darwin, ‘‘ Fertilization of Orchids,” 2nd edit., pp. 191, 192.

REFLEX ACTION 31

consciousness. Among plants we have many interesting and
admirable examples of organic behaviour; but nowhere so
much as a hint of that profiting by individual experience

Fic. 10.—Catasetum ; OC, diagram of column; a, anther; an, horn; d, ad-
hesive disc; f, filament of anther; g, ovarium; ped, pedicel; D and
E, pollinium ; p, pollen-mass. (From Darwin’s “ Orchids.” )

which is the criterion of the effective presence of conscious
guidance and control.

V.—REFLEX ACTION

It is sometimes said that the tentacles of the Sun-dew leaf
indicate a primitive kind of reflex action in plants, and that
they afford evidence of discrimination. “It is,” says Romanes,

32 ORGANIC BEHAVIOUR

“the stimulus supplied by continuous pressure that is so
delicately perceived, while the stimulus supplied by «mpact is
disregarded.” * And, comparing this with what is observed
in the Venus’s Fly Trap, ke says: “In these two plants the
power of discriminating between these two kinds of stimuli
has been developed to an equally astonishing extent, but in
opposite directions.” | It is well, however, to avoid terms
which carry with them so distinctively a conscious implication -
as “ discrimination ” and “ perception ” do for most of us. Just_
as the photographer’s film reacts differently according to the
quality of light-rays, violet or red, which reach it, so do many
organic substances react differently to stimuli of different
quality, irrespective of their intensity. The “ discrimination ”
of plants and of some of the lower animals is of this kind,
and it is better to speak of it simply as differential reaction.
There can then be no chance of its being confused with con-
scious choice.

Nor should the movements of the Sun-dew tentacles or of
those of the Sea-anemone be termed in strictness reflex action.
As originally employed by Marshall Hall, and, since that time,
by common consent, reflex action involves a differentiated
nervous system. There is, first, an afferent impulse from -
the point of stimulation passing inwards to a nerve-centre ;
secondly, certain little-understood changes within this centre ;
and thirdly, an efferent impulse from the centre to some organ
or group of cells which are thus affected. In plants there is
no indication of anything analogous to this specialized mode
of response. The impulse passes directly from the point of
stimulation to the part affected without the intervention of
anything like a nerve-centre. In the sensitive Oxalis the
impulse passes directly to the point of insertion of the leaflet
or leaf-stalk ; in Catasetum, from the horn to the retaining
membrane; in the Sun-dew, from the affected tentacle to
those in its neighbourhood. Even in the Sea-anemone,
though there is a loosely diffused nervous system, the passage

* “Mental Evolution in Animals,” p. 50.
t Ibid., p. 51.

REFLEX ACTION 33

of the impulse from one part of the circlet of tentacles to
other parts, seems to follow a direct rather than a reflex
zourse, and there do not appear to be any specialized centres
by which the impulses are received and then redistributed.

In all animals in which well-differentiated nervous systems
are found, in which there are distinct nerve-fibres and nerve-
centres, reflex actions, simple or more complicated, occur.
They form the initial steps leading up to the highest types
of organic behaviour. So long as the nervous arcs—afferent
fibres, nerve-centre, and efferent fibres—remain intact reflex
acts may be carried out with great precision and delicacy,
even when the higher centres, which we believe to be those of
conscious guidance and control, have been destroyed. When,
for example, the whole of the brain of a frog has been extir-
pated and the animal is hung up by the lower jaw, if the left
side be touched with a drop of acid the left lee is drawn up
and begins to scratch at the irritated spot, and when this leg
is held, the other hind leg is, with seemingly greater difficulty,
brought to bear on the same spot. ‘ This,” says Sir Michael
Foster, “at first sight looks like an intelligent choice... .
But a frog deprived of its brain so that the spinal cord only
is left, makes no spontaneous movements at all. Such an
entire absence of spontaneity is wholly inconsistent with the
possession of intelligence. ... We are therefore led to con-
clude that the phenomena must be explained in some other
way than by being referred to the working of an intelli-
gence.” * But if we concede that intelligence is absent, may
there not at least be some consciousness? Sir Michael
Foster’s reply to such a question goes as far as we have any
justification for going, even when we give free rein to conjec-
ture. “We may distinguish,” he says, “ between an active
continuous consciousness, such as we usually understand by
the term, and a passing or momentary condition, which we
may speak of as consciousness, but which is wholly discon-
tinuous from an antecedent or from a subsequent similar
momentary condition; and indeed we may suppose that the

* «A Text-book of Physiology,” 5th edit., part iii., p. 909.
D

34 ORGANIC BEHAVIOUR

complete consciousness of ourselves, and the similarly com-
plete consciousness which we infer to exist in many animals,
has been evolved out of such a rudimentary consciousness.
We may, on this view, suppose that every nervous action of a
certain intensity or character is accompanied by some amount
of consciousness which we may, in a way, compare to the light
emitted when a combustion previously giving rise to invisible
heat waxes fiercer. We may thus infer that when the brain-
less frog is stirred by some stimulus to a reflex act, the spinal
cord is lit up by a momentary flash of consciousness coming
out of the darkness and dying away into darkness again ; and
we may perhaps infer that such a passing consciousness is the
better developed the larger the portion of the cord involved
in the reflex act and the more complex the movement. But
such a momentary flash, even if we admit its existence, is
something very different from consciousness as ordinarily
understood, is far removed from intelligence, and cannot be
appealed to as explaining the ‘ choice’ spoken of above.” *
These sentences indicate with sufficient clearness the
distinction, more than once hinted at in the foregoing pages,
between consciousness as an accompaniment, and conscious-
ness as a guiding influence. We shall have more to say in
this connection in subsequent chapters. The experiment with
the frog shows, at any rate, that reflex actions, of a distinctly
purposive nature, may be carried out when the centres, which
are believed to exercise conscious control and guidance have
been destroyed. It is said that in man, when, owing to
injuries of the spine, the connection between the brain and
the lower part of the spinal cord have been severed, tickling
of the foot causes withdrawal of the limb without directly
affecting the consciousness of the patient. Bunt in all such
cases we are dealing with a maimed creature. The living frog
or man, healthy and intact, is, presumably in the one case,
certainly in the other, conscious of these reflex actions, and
can exercise some amount of guidance and control over them.
In man this is unquestionably the case. But granting that

* “A Text-book on Physiology,” 5th edit., part iii., pp. 911, 912.

[THE EVOLUTION OF ORGANIC BEHAVIOUR 35

he brain is the organ of conscious control, granting that it
‘an receive impulses from and transmit impulses to the reflex
‘entres, no more is here implied, and no more can be legiti-
nately inferred, than that the kind of organic behaviour we
‘all “reflex action” is in the higher animals in touch with
the guiding centres. We have no ground for assuming that
n reflex action there is any power of intelligent guidance
ndependent of that which is exercised by the brain or
malogous organ. In brief, reflex acts, in animals endowed
vith intelligence, may be regarded as specialized modes of
wrganic behaviour; which are in themselves often charac-
verized by much complexity ; which subserve definite biological
mds; which are effected by subordinate centres capable of
ransmitting impulses to, and receiving impulses from, the
sentres of intelligent guidance; and which, as responses con-
ined to certain organs or parts of the body, form elements
in the wider behaviour of the animal as a whole.

VI.—THE EvoLUutTion OF ORGANIC BEHAVIOUR

The interpretation of organic behaviour in terms of evolu-
sion mainly depends on the answer we give to the question :
Are acquired modes of behaviour inherited? A negative
uswer to this question is here provisionally accepted. But
ihe premisses from which this conclusion is drawn are too
sechnical for discussion in these pages. It must suffice to
state as briefly as possible what this conclusion amounts to,
md to indicate some of the consequences which follow from
ts acceptance.

The fertilized egg gives origin, as we have seen, to the
nultitude of cells which build up the body of one of the
nigher animals. There are, on the one hand, muscle-cells,
rland-cells, nerve-cells, and other constituents of the various
issues; and there are, on the other hand, the reproductive
ells—ova or sperms, as the case may be. Now, every cell in
hhe developed animal is a direct descendant of the fertilized
‘gg. But of all the varied host only the reproductive cells

36 - ORGANIC BEHAVIOUR

take any direct share in the continuity of the race. Here-
ditary transmission is therefore restricted to the germinal
substance of these reproductive cells. Trace the ancestry of
any cell in the adult body, say a nerve-cell, and you reach the
fertilized ovum. Trace back the ancestral line yet further,
and you follow a long sequence of reproductive cells, or, at
least, of cells which have undergone but little differentiation ;
but never again will you find, in the course of a genealogy of
bewildering length, a nerve-cell. Such a tissue-element is a
descendant, but cannot. become an ancestor; it dies without
direct heirs.

It is universally admitted that the bodily structures are
subject to what is termed modification under the stress of
environing circumstances. The muscles may acquire unusual
strength by use and exercise; the nerve-centres may learn
certain tricks of behaviour in the course of individual life ; and
other structures may be similarly accommodated to the con-
ditions which affect them. To such modifications of structure
or function in the organs or parts the term acquired is
primarily applied. The tissues have thus a certain amount
of organic plasticity, through which they are adjusted to a
range of circumstances varying in extent. They are able to
acquire new modes of behaviour. But the cells of which they
are composed are off the line of racial descent. They leave
no direct heirs. When the body dies the modifications of
behaviour acquired by its parts perish with it. Only if in
some way they exercise what we may term a homceopathic
influence on the germinal substance can the accommodation
they have learnt be transmitted in inheritance. By a homeo-
pathic influence is here meant one that is of such a nature as
to communicate to the germinal substance, the seeds of similar
changes of structure or function. And of the occurrence of
any such homeopathic influence there is no convincing
evidence.

Logically contrasted with the modifications of the tissues,
dependent on organic plasticity, are the variations which arise
from the nature and constitution of the reproductive cells.

THE EVOLUTION OF ORGANIC BEHAVIOUR 37

How they arise cannot here be discussed. But they are, it is
believed, subject to the influence of natural selection, which
has guided them, throughout the ages of organic evolution,
in the directions they have taken ; disadvantageous variations
having been eliminated, and favourable variations surviving
in the struggle for existence. Such modes of behaviour as are
congenital and are due to hereditary transmission are therefore
the outcome of variations which have been selected generation
after generation. And the fit adjustment of this congenital
behaviour to the needs of life is termed adaptation. It is here
assumed that modifications of behaviour in one generation
are not inherited, and therefore contribute nothing to the
store of adaptive behaviour in the next generation.

It must not, however, be supposed that the provisional accept-
ance of this conclusion involves the denial of all connection
of any sort between accommodation and adaptation. When we
remember that plastic modification and germinal variation have
been working together, in close association, all along the line of
organic evolution to reach the common goal of adjustment to
the circumstances of life, it is difficult to believe that they
have been throughout the whole process altogether independent
of each other. Granted that acquired modifications, as such,
are not directly inherited, they may none the less afford the
conditions under which coincident variations escape elimination.
By coincident variations I mean those the direction of which
coincides with that taken by modification. The survival of
an animal depends on its adjustment to the circumstances
of its life, no matter how this adjustment is secured. And
this survival would in the long run be better secured, we may’
suppose, where the two methods of adjustment were coincident
and not conflicting ;* just as a man who not only acquires by

* Professor Mark Baldwin has applied the term “organic selection ”
to the result of this interaction (American Naturalist for June and
July, 1896). Cf. also H. F. Osborn (Sedence, Nov. 27, 1896); August
Weismann (Romanes Lecture on “The Effects of External Influences
on Development,” 1894), and “Germinal Selection,’ Monist, Jan.,
1896; and the author’s “ Habit and Instinct,” ch. xiv., 1896.

38 ORGANIC BEHAVIOUR

his own exertions a fortune but also inherits one, is better off
than his neighbour, of equal business capacity, who is entirely
dependent on his own exertions. The inheritance of a small
capital may, indeed, make just the difference between success
and failure. Even with it, if he had no power of acquiring
more, he might remain a poor man. Inheritance and acqui-
sition combined may best lead to survival in competition.
Thus modification may supply the conditions under which
coincident variations are favoured, and, given time, to reach
step by step, through natural selection, a fully adaptive level.
If this be so we may accept many of the facts adduced by the
transmissionist in favour of the direct inheritance of acquired
characters, and at the same time interpret them on selectionist
principles.

If, however, acquired characters are not hereditary the
method of natural selection in racial progress is curiously
indirect. Apart from the preservation of their fecundity, the
cells on which the continuity of life, in all the higher animals,
depends, have themselves taken little part in the struggle for
existence. Just as in the forest tree, the firmly implanted
roots, the sturdy stem, and the strong branches have to bear
the stress of the winter storm, that the flowers of spring may
ripen the seeds which contain the potentiality of all this
strength ; so do muscle, sinew, and brain secure the survival
of the animal, that his descendants may carry on the struggle.
One may liken the cellular constituents of the animal to a hive
of bees with fertile drones and queen, and sterile workers. It
is on the exertions of the latter that, in the struggle for
existence, the continued existence of the swarm depends, while
it is by the pairing of the fertile drone and queen that the
continuity of the race is secured. No worker can transmit
the qualities which are so essential to the well-being of the
community. But in the eggs of their sister the queen-mother
these qualities lie dormant. And since the race is one race,
the workers by their exertions contribute indirectly to the
maintenance of those hereditary aptitudes to which they are
unable to contribute directly. For it is essential to bear in

THE EVOLUTION OF ORGANIC BEHAVIOUR 39

mind that they not only work for their own generation, but
they determine the course of heredity. Picture two such
communities set in an environment which intensifies the
struggle for existence. The one is strong, healthy, and
vigorous ; the other in all respects the reverse. The incidence
of the battle of life falls mainly on the workers. If they
succumb in the one group their fertile queen either perishes,
or gives rise to a poor stock, certain in the Jong run to be
eliminated. But the vigorous workers in the other group
survive and secure, too, the survival of their queen, who, since
she is also their sister, bears, in her ovaries, the good seed from
which a new generation of vigorous workers shall be developed.
Thus though the sterile bees contribute nothing directly to
the heredity of the race, they indirectly determine the direction
which that heredity shall take. So, too, in the higher animals,
the reproductive cells are the fertile sisters of a host of sterile
body cells, on which the main incidence of the struggle for
existence falls. Their sterility precludes their directly con-
tributing to the success of future cell-generations ; but in
protecting their fertile sisters, the reproductive cells, they are
really determining the lines along which the evolution of the
race shall continue.

Acquired characters may thus be regarded as the results of
those accidents, fortunate or the reverse as the case may be,
which happen to the body, and more or less modify its outward
form or hidden structure, and its modes of organic behaviour ;
but which, as such, have no direct effects for better or worse
on the germinal substance. All that the plant or animal can be
is due to heredity ; all that it és, to heredity and circumstance.
Even the ability to yield to circumstance is part of heredity’s
dower. Fortunate, then, the plant or animal that inherits such
definiteness of structure and behaviour as may fit it to its
station, together with such plasticity as may enable it to
accommodate itself to those changes of environing conditions
which may fall to its lot.

One more point must be noticed in connection with
this difficult and puzzling subject. The acceptance of the

40 ORGANIC BEHAVIOUR

conclusion that acquired modes of behaviour are not here-
ditary nowise commits us to the belief that heredity has nothing
whatever to do with them. Though what is acquired may
not be transmitted, what one may term the acquisitiveness is
unquestionably inherited. Though this, that, or the other
acquired mode of behaviour may have no direct descendants,
the power of acquiring any one of them under the appropriate
circumstances is handed on as an invaluable legacy. Just as
the mirror which has reflected a fleeting scene retains no
lasting image of the bygone events, so heredity may retain no
impress of acquired characters; but just as the mirror keeps
its power of reflecting such scenes, so does heredity transmit
the power of acquiring such characters. As the leaves of the
oak are renewed each successive spring, so may acquired modes
of behaviour be repeated in each successive generation if only
the requisite conditions recur in due season.

From what has preceded it may, therefore, be inferred that
organic behaviour may arise either through modifications
occurring in the plastic tissues, or through variations having
their origin in the germinal substance. Broadly speaking,
however, we may regard as predominantly due to adaptation
those congenital modes of behaviour and those organic re-
sponses which on their first occurrence are relatively definite
in character, and which are directed to a biological end, for
whose attainment the tissues have had no preparatory training ;
and we may regard as predominantly due to accommodation
those responses which are, so to speak, learnt by the tissues in
the course of individual life. Both are dependent on heredity,
but in different ways. What the animal owes to heredity may,
indeed, as I have elsewhere said,* be classified under two
heads. Under the first will fall those relatively definite modes
of behaviour which fit the animal to deal at once, on their
first occurrence, with certain essential or frequently recurring
conditions of the environment. Under the second head will
fall the power of dealing with special circumstances as they
arise in the course of a varied life. The former may be

* “ Habit and Instinct,” p. 26.

ara alt ee”

THE EVOLUTION OF ORGANIC BEHAVIOUR 41

likened to the inheritance of specific drafts for definite needs
which are sure to arise in the conduct of life; the latter
to the inheritance of a legacy which may be drawn upon for
any purpose as occasion may demand. If the need becomes
habitual the animal may, so to speak, instruct his banker to set
aside a specific sum to meet it as it arises. But this arrange-
ment is a purely individual matter, dictated by experience, and
in no wise enjoined by the original terms of the bequest. And
both types are fostered by natural selection which develops
(#) such congenital definiteness of response, and (0) such
innate plasticity, as are advantageous under the conditions
of existence ; uniform conditions tending to emphasize the
former, variable conditions the latter.

Difficult as it may be to earmark the items of the organic
bequest—to say that, of the sum of energy expended in any
given case of organic behaviour, so much is due to a specific
draft definitely assigned in heredity for this particular purpose,
and so much is contributed from the general legacy of innate
plasticity,—it none the less conduces to clear thinking to empha-
size the logical distinction between them, so long as it is
steadily borne in mind that logical distinction does not imply
biological separation. The animal, with all its varied modes
of behaviour, is an organic whole, and as an organic whole it
has been developed from the fertilized egg. The very same
tissues which exhibit congenital modes of behaviour are
capable also of acquiring new responses and playing their part
in accommodation. We have not one set of organs which are
the products of variation and another set which result from
modification. Our study would no doubt be simplified if this
were the case ; but it is not so. And we must take the animal
as we find it, presenting varied behaviour of complex origin.
Even the reflex nervous centres, which are concerned in
responses so automatic as to suggest a stereotyped structure of
distinctively germinal origin, are also, as we saw at the close of
the last section, in close touch with those centres of control
which are associated with the supreme power of accommoda-
tion arising from the possession of consciousness.

CHAPTER II
CONSCIOUSNESS

I.—TueE Conscious ACCOMPANIMENTS OF CERTAIN ORGANIC
CHANGES

Ir is possible that all organic behaviour is accompanied by
consciousness. But there is no direct means of ascertaining
whether it is so or not. This is, and must remain, a matter of
more or less plausible conjecture. We have, indeed, no direct
knowledge of any consciousness save our own. Undue stress
should not, however, be laid on this fundamental isolation of
the individual mind. We confidently infer that our fellow-
men are conscious, because they are in all essential respects
like us, and because they behave just as we do when we act
under its guiding influence. And on similar grounds we
believe not less confidently that many animals are also con-
scious. But how far we are justified in extending this
inference it is difficult to say. Probably our safest criterion
is afforded by circumstantial evidence that the animal in
question profits by experience. If, as we watch any given
creature during its life-history, we see at first a number of
congenital or acquired modes of behaviour, we may not be
able to say whether they are accompanied by consciousness or
not ; but if we find that some of these are subsequently carried
out more vigorously while others are checked, we seem justified
in the inference that pleasurable consciousness was associated
with the results of the former, and disagreeable consciousness
with those of the latter. When we see that a chick, for
example, pecks at first at any small object, it is difficult to
: 42

AS ACCOMPANIMENT OF ORGANIC CHANGES 43

say, on these grounds, whether it is a sentient animal or only
an unconscious automaton ; and if it continued to behave in
a similar fashion throughout life, our difficulty would still
remain. But when we see that some objects are rejected
while others are selected, we infer that consciousness in some
way guides its behaviour. The chick has profited by experience.
But even this is clearly only a criterion of what we may term
effective consciousness. There may be sentience which is
merely an accompaniment of organic action without any
guiding influence on subsequent modes of behaviour. In that
case it is not effective; and whether it is present or not we
have no means of ascertaining.

We seem also to be led to the conclusion, both from a
priort considerations and from the results of observation, that
effective consciousness is associated with a nervous system.
Its fundamental characteristic is control over the actions, so
that some kinds of behaviour may be carried out with increased
vigour, and others checked. And it is difficult to see how this
can take place unless the centres of control are different from
those over which they exercise this influence. If we are to
understand anything definite by the guidance of consciousness,
we must conceive it as standing apart from and exercising an
overruling influence over that which it guides. This is un-
questionably an essential characteristic of consciousness, as
generally understood by those who take the trouble to con-
sider its relation to behaviour ; and though some would seek
to persuade us that a mere accompaniment of consciousness
can somehow determine the continuance or discontinuance of
organic behaviour, it is difficult to see how this can be the
case. The accompaniment of air-tremors can no more influence
the vibrations of a sounding string than an accompaniment of
consciousness can affect the nature of the organic changes in
the tentacles of the Sun-dew leaf.

And if, instead of trusting to such general « priori con-
siderations, we study with attention the conditions under which
an animal so behaves as to lead us to infer that it profits
by experience, we find that it is not the consciousness that

44° CONSCIOUSNESS

accompanies the behaviour which leads to future guidance, but
the consciousness that arises from the results of the behaviour.
Let us willingly grant that the newly hatched, and as yet in-
experienced chick, when it pecks at a small object is conscious
of a visual impression, and is conscious also of movements of
its neck and beak. These do not constitute the experience by
which it profits. This is provided by the results of the peck-
ing, according as the morsel seized is nice or nasty. We may
say, in popular language, that the little bird remembers when
it sees a similar object that the former results were pleasant or
distasteful, as the case may be; and that it is through this
remembrance that future guidance is rendered possible. But
all the evidence that we possess goes to show that the sensory
centres, stimulated by what we will assume to be the taste of
the morsel, are different from those which are affected by
sight, and the movements concerned in pecking. So that the
consciousness which is effective in guiding future action is an
accompaniment of the stimulation of centres that are different
from those concerned in the behaviour over which guidance
is exercised. And if this interpretation of the observed facts
be correct, it supports the conclusions reached from @ priori
considerations. It seems further to show that, not only is a
nervous system necessary for the occurrence of controlled
behaviour, but that no little complexity in its intercommunica-
tions is essential.

It may be urged that the chick’s behaviour which has
been selected for purposes of illustration, and the inter-
pretation we have put upon it, throws too much stress on
remembrance, so called, and further gives the false impression
that all experience must be for future guidance. There are
surely numberless cases, it will be said, in which nothing of
the nature of distinct memory is involved, and in which the
guidance of consciousness is exercised at once over present
behaviour, without any postponement to the future. Even
omitting for the present the former point, the formula implied
—that present experience is for future guidance—cannot be
accepted in view of the familiar fact that present experience is

AS ACCOMPANIMENT OF ORGANIC CHANGES 4s

constantly influencing present behaviour. Practically speak-
ing, this is perfectly true: because, practically, under the
term present we include quite an appreciable period of time—
say, a few seconds, or even minutes. If we narrow our con-
ception of the present, as is commonly done in philosophical
discussions, to the boundary line between past and future, then
it will be seen that even the guidance of what in popular
speech is called present behaviour is really exercised on the
subsequent phases of that behaviour. At the risk of some
technicality our position must be explained a little more fully.
It is assumed that the data of consciousness are afforded by
afferent impulses coursing inwards from the organs of special
sense, or those concerned in responsive movements. This
conclusion rests on such a wide body of psychological inference
that it may be accepted without discussion, at any rate for
our immediate purpose. The efferent impulses, those which
effect the orderly contraction of the muscles, are unconscious ;
but when the movement is produced afferent impulses course
inwards from the parts concerned in the behaviour, and these
may then contribute data to consciousness. ,
Now let us suppose that a chick, which has been hatched in
an incubator, be removed some twelve hours after birth, held
in the hands for a few minutes until its eyes have grown
accustomed to the light, and placed on a table near some small
pieces of hard-boiled egg. Let us watch its behaviour and
endeavour to interpret it. We shall have occasion to consider
hereafter whether the conscious experience of parents and
ancestors is inherited as such ; for the present we will assume
that it is not. The chick has to acquire for itself its own
experience. A piece of egg catches the eye of the little bird,
which then pecks at it, and just fails to seize it. Here is a piece
of congenital organic behaviour. Taken by itself one might
find it difficult to say whether it is accompanied by conscious-
ness or not, just as one finds it difficult to say whether the
closure of the Venus’s Fly-trap is conscious. But the sub-
sequent behaviour of the chick leads us to infer that it is a
sentient animal ; and we may, therefore, fairly assume that it

46 CONSCIOUSNESS

is sentient from the first. Dividing the course of the observed
behaviour into stages, we may say that the first stage is that in
which the chick receives a visual stimulus accompanied by a
sensation of sight. Upon this there rapidly follows the second
stage, when the bird pecks, and its experience is widened by
new data of consciousness derived from a group of motor
sensations ; and upon this, again, there follows the third stage,
when sensations come in from the morsel of egg which the
chick touched but just failed to seize. After a pause the chick
strikes again. But we have not a mere repetition of the
former sequence of stages. The visual stimulus at first fell
upon the eye of a wholly inexperienced bird ; now it falls upon
the eye of one that has gained experience of pecking and
tasting. What we may call the conscious situation has com-
pletely changed, at all events if we assume that the items of
consciousness, including as essential the consciousness of be-
haviour, do not remain separate and isolated, but have coalesced
into a group through association. And in this group the
consciousness of behaving is perhaps the most important
element in the situation, making it of practical value. What
psychologists term the presentative visual stimulus, now calls up
re-presentative elements, motor and gustatory ; and these place
the situation in a wholly new aspect. They give it what Dr.
Stout terms “meaning.” On the second or third attempt
the chick seizes and swallows the morsel of egg. Its ex-
perience is yet further widened ; and thereafter the situation
has other new elements. Later it pecks at some nasty grub ;
shakes its head, and wipes its bill on the ground. The con-
scious situation has for the future become more complex,
and the behaviour is henceforth differentiated into that of
acceptance and that of rejection, in each case determined by
the acquired meaning in the coalescent conscious situation :
the sight of a nice piece of egg being one situation, that of
a nasty caterpillar another, each associated with its specific
behaviour-consciousness. We need not carry the illustration
further on these lines: the essential feature is that experience
grows by the coalescence of successive increments, and that

AS ACCOMPANIMENT OF ORGANIC CHANGES 47

each increment modifies the situation which takes effect on the
succeeding phases of behaviour, even if they succeed within the
fraction of a second. That is what is meant by saying that
present experience is for future guidance. The future need
not be remote, but may be so immediate that in popular speech
we may say that it is not future but present guidance which is
rendered possible.

We may now turn for a moment to the criticism that there
are numberless cases in which nothing of the nature of distinct
memory is involved. We may now substitute for the word
remembrance, which was used above, the more technical term
re-presentation. Profiting by experience, regarded as a criterion
of the presence of effective consciousness, involves re-presenta-
tive elements in the conscious situation which carry with them
meaning. Let us for the moment assume an ultra-sceptical
attitude with regard to any conscious accompaniment. The
chick when it pecks, let us say, is an unconscious automaton.
It seizes a piece of egg; this affords an unconscious stimulus,
which sets agoing unconscious acts of swallowing ; or it seizes
a piece of meal soaked in quinine, which sets agoing unconscious
acts of rejection and touches the hidden springs which make
the automaton wipe its bill. So far we find no great difficulty.
It is when we have to consider subsequent behaviour that
a severe strain is felt on this method of interpretation.
One can understand an automatic action repeated again and
again as often as the stimulus is repeated. But the chick
may shake its head and wipe its bill on the mere sight of
the quinine-soaked meal, which, on the hypothesis of conscious
experience, has already proved distasteful. So that if we
accept the unconscious automaton theory we must assume an
organic association which closely simulates the conscious
association to which our own experience testifies. But the
associations which take part in the guidance of behaviour in
the chick are so varied and delicate, so closely resemble those
which in ourselves imply conscious guidance, that a sceptical
attitude throws more strain upon our credulity than the
acceptance of the current belief in conscious control. We

48 CONSCIOUSNESS

shall therefore assume that evidence for such coalescent
association is also evidence of the presence of effective
consciousness.

Tt may still be said, however, that in selecting an example
from so highly organized an animal as a bird, we are taking
for granted that a complex case of controlled behaviour may
fairly be accepted as a type of more simple cases. Unfortunately
the only being with whose power of conscious control we have
any. first-hand acquaintance is possessed of a nervous system
even more complex than that of the chick. Our psychological
interpretations are inevitably anthropomorphic. All we can
hope to do is to reduce our anthropomorphic conclusions to
their simplest expression. The irreducible residuum seems to
be that wherever an animal, no matter how lowly its station in
the scale of life, profits by experience, and gives evidence of
association, it must have some dim remembrance, or, let us now
say, Some re-presentation, of the results of previous behaviour
which enters into and remodels the conscious situation ; that
through the re-presentative elements behaviour is somehow
guided ; and, further, that the centre of conscious control is
different from the centre of response over which the control is
exercised.

Il.—Txe Earty Stages or Mentat DEVELOPMENT

We use the phrase “ mental development” in its broadest
acceptation as inclusive of, and applicable to, all phases of
effective consciousness. We shall assume that throughout
this development there is a concomitant development of nerve-
centres and of their organic connections. And we shall further
assume that experience, as such, is not inherited.

The nature of the grounds on which the latter assumption
is based must first be briefly indicated. It is commonly
asserted that fear of man, the inveterate hunter and sportsman,
is inherited by many animals, as is also that of other natural
enemies. This is, however, questioned, or even denied, by
many careful observers. Mr, W. H. Hudson has an excellent

EARLY STAGES OF MENTAL DEVELOPMENT 49

chapter on “ Fear in Birds” in his ‘ Naturalist in La Plata,”
and concludes that: fear of particular enemies is, in nearly all
cases, the result of experience individually acquired. I have
found that pheasants, partridges, plovers, domestic chicks, and
other young birds, hatched in an incubator, show no signs of
fear in the presence of dog or cat, so long as the animal is not
aggressive. It should be mentioned, however, that Miss M.
Hunt * asserts that chicks do show inherited fear of the cat. Dr.
Thorndike’s t observations, on the other hand, support my own,
which I have since repeated with the same results. Neither
birds nor small mammals show any signs of fear of stealthily
moving snakes. My fox terrier smelt, nose to nose, a young
lamb which was lying alone ina field. I was close at hand, and
could detect no indication of alarm on the part of the lamb till
the mother came running up in great excitement. Then the
lamb ran off to her dam. Whenever opportunity has arisen,
I have introduced young kittens to my fox terrier, and have
never seen any sign of inherited fear. He was a great hunter
of strange cats, but was trained to behave politely to all
birds and beasts within the precincts of my study. It is true
that he was on good terms, or at least terms of permissive
neutrality, with the kittens’ mother. And it may be said that
this was inherited ; but such an argument cannot apply in the
case of pheasant or lamb.

Here, as throughout our study of animal behaviour in its
conscious aspect, we have not only to conduct observations
with due care, but to draw inferences with due caution.
Douglas Spalding described how newly hatched turkeys
showed signs of alarm at the cry of a hawk ; and he inferred
that, since this sound was quite new to their individual expe-
rience, the alarm was due to the inheritance of ancestral
experience of hawks. But since young birds show signs of
alarm at any sudden and unaccustomed sound—a sneeze, the
noise of a toy horn, a loud violin note, and so forth—the safer
inference seems to be that they may be frightened by strange

* American Journal of Psychology, vol. ix., No. 1.
+ Psychological Review, vol. vi, No. 3.

5° CONSCIOUSNESS

sounds of many kinds. But this does not imply the inheri-
tance of experience, which is essentially a discriminating pro-
cess. There is no sufficient evidence that a peculiar cry
suggests the hawk, of which the progenitors have acquired
bitter experience ; nothing to justify the belief that the sound
carries with it inherited meaning. And as with hearing, so
with sight. Young birds may be frightened by many strange
objects. I have seen a group of several species, filled with
apparent alarm at a large white jug suddenly placed among
them, at balls of paper tossed towards them, at a handkerchief
dropped in their midst. Itis, in fact, their inexperience which
is often the condition of such fear. As Mr. Hudson says :*
“A piece of newspaper carried accidentally by the wind is as
great an object of terror to an inexperienced young bird as a
buzzard sweeping down with death in its talons.”

Until recently it was commonly asserted that birds avoid
gaudy but nauseous or harmful insects through the inheri-
tance of experience gained by their ancestors through many
generations. But here again the inference seems to have been
incautiously drawn. Of the hundreds of young birds I have
had under observation, not one has avoided the peculiarly dis-
tasteful cinnabar caterpillar, until it had gained for itself
experience of its nauseous character. So too of wasps and
bees. Only through experience are these avoided. It is true
that chicks may shrink from them if they buzz or even walk
rapidly towards them. But a large harmless fly will inspire
just as much timidity. As the result of careful observations,
Mr. Frank Finn t concludes “that each bird has to separately
acquire its experience, and well remembers what it has learnt.”
And with this conclusion my own observations are entirely in
accord.

Such is some of the observational evidence on which is
based the provisional hypothesis that experience, as such, is
not inherited. What, then, is inherited? Clearly the organic
conditions under which experience can be acquired. Since a

* “Naturalist in La Plata,” p. 88,
+ Journal Asiatic Society of Bengal, \xvii., part ii., 1897, p. 614.

EARLY STAGES OF MENTAL DEVELOPMENT | 51

young bird inherits a tendency to peck at small objects, espe-
cially, in the case of some birds such as plovers or partridges,
at small moving objects, opportunities are afforded for dis-
crimination in accordance with the results of experience.
Since its inherited timidity leads the chick to shrink from
many things seen or heard, a wide range of conscious data is
supplied. Inheritance provides the raw material of organic
behaviour for effective consciousness to deal with in accord-
ance with the results which are its data.

Having thus cleared the ground and laid bare some at
least of the assumptions which we accept as foundations on
which to build, we may now follow up the, line of treatment
which was suggested in the first section of this chapter.
Remembering that our aim is to understand the influence of
consciousness on behaviour—or, in more accurate, if more
cumbrous phraseology, the influence of certain nerve-centres
which have for their concomitant what we have termed effec-
tive consciousness—the questions which present themselves
in any given case are: What is the conscious situation which
is effective in guidance ? what elements enter into the situa-
tion, whence are they derived, and how were they introduced ?
how do they take effect in behaviour ?

If it be true that, in many of the lower forms of life, con-
sciousness or sentience, though presumably present in some
dim form, is merely an accompaniment of organic behaviour
without reaching the level of recognizable effectiveness ; and
if, during the development of one of the higher animals from
the fertilized ovum, the early stages of organic behaviour are
in -like manner merely sentient ; it follows that, when effec-
tive consciousness enters upon the scene (who can say at
what exact stage of evolution ?), it finds itself a partner in a
going concern. Much organic business is being transacted
with orderly regularity ; preparations have been made for
more extensive operations; and energies lying dormant, or
expending themselves aimlessly in starts and twitches, await
the guidance which shall direct them to higher and wider
biological ends. Or, to vary the analogy, consciousness is

52 CONSCIOUSNESS

the heir to a wide estate, over which he has no control until
he comes of age. Up to that time the estate is managed in
strict accordance with the dictates of the hereditary bequest.
He may be aware of what is going on, but merely as a spec-
tator without power of interference. And when he comes into
possession his first business is to gather up the threads. He
must learn bit by bit how the estate is being managed, that he
may have data for the guidance of his own management
within the wise limits of the hereditary entail.

Now, when a mammal is born, a bird is hatched, an insect
emerges from the chrysalis, we have, if not the beginning, at
any rate a great and sudden extension of the range of effective
consciousness. In the case of the mammal and bird the
experience gained in the womb or within the egg-shell is
presumably of little value for the wider life upon which an
entrance is made. It is true that an insect has passed through
a previous stage of active and no doubt consciously guided
existence as a caterpillar. But we do not know whether the
experience thus acquired is effectual for use in the later imago
stage. And we may perhaps infer from the extensive re-
modelling of the nervous system, which occurs during the
chrysalis sleep, that this itself serves to break the continuity
of experience. In any case the newly hatched chick, if it
inherit no experience, and can have gained little of guiding
value in the egg, enters upon a situation which from the
number and variety of the data supplied may well seem to
‘us bewildering. If we picture ourselves in such a position,
with sights, sounds, motor sensations, touches, and pressures
raining in upon a virgin experience, we wonder how we should
make a beginning; how we could possibly decide on the first
step towards reducing this multiplicity and. diversity to some-
thing like unity and order. And perhaps we wonder how we
ourselves made a beginning when we were pink newly born
babies.

If it may be said without paradox, we never did make a
beginning. The beginning was made for us. For we habitually
associate ourselves with the control centres, and regard our

EARLY STAGES OF MENTAL DEVELOPMENT 53

bodies, like our watches, as ours and not us. We wind the
bodily watch, and set its hands from time to time; but we
did not make it, and it was already going when heredity
handed it to us over the counter of birth. The first step
towards reducing the seeming chaos of sensory experience to
something like order is not due to the selection by conscious-
ness of this or that element for prominence among the rest,
but to the thrusting forward of certain modes of behaviour:
by the conditions of organic life. The differentiation of the
field of vision in the chick is not effected by any conscious
determination to fix the attention on that wriggling maggot,
but through the congenital response it calls forth. This serves
not only to make the grub stand out clearly amid its surround-
ings, but also to emphasize a motor group, called into vigorous
action in the midst of other motor ‘sensations, and, in rapid
sequence, to lay stress on a sensation of taste suddenly called
into prominence.

Nor, as we have seen, do the organic effects cease here.
The functional action of three sensory centres is thus called
into play. But they are constituent parts of one nervous system.
The direct stimulation of each by nerve impulses from eye,
motor organs, and beak, gives temporary predominance to
certain sensory data which are termed presentative. But the
several centres are connected with each other. And thence-
forward, in subsequent stages of experience, the direct stimula-
tion of the visual centre indirectly calls into play the other
two, so that the presentation through sight evolves re-presenta-
tions of the motor group, and of taste. Hence sentience is
not sufficient for guidance ; there must be consentience involving
the presence of several elements. But these elements must not
be regarded as separate save in our analysis ; they form con-
stituent parts of the coalescent situation as a whole, of which
alone the chick is presumably conscious, without analysis of
detail.

It is just because the chick is a going concern when con-
sciousness comes of age and begins to assume control—just
because a wide range of congenital behaviour is part of the

54 CONSCIOUSNESS

organic heritage—that the early stages of the acquisition of
experience proceed so rapidly and so smoothly. The animal
has not to make and fashion the early conscious situations ;
it has only to accept them. It has not at first to enforce
order on the multiplicity of sensory data raining in upon the
conscious centres; it has only to take note of the existing
order among them. It has not painfully to learn how to
co-ordinate the efferent impulses proceeding to the many
muscles concerned in some simple response; it has only to be
sensitive to the response as a whole. It has not to select the
association of this, that, and the other group of data within a
coalescent situation ; organic behaviour provides it with pre-
determined sequences ready made—sequences which have for
generations received the emphatic sanction of natural selection.
Congenital tendencies which it has inherited but not acquired
determine all its earliest behaviour, determine what elements
in the sensory complex shall be thrust into conscious pro-
minence, determine in what manner these data shall be asso-
ciated ; determine, in fact, what salient points in the developing
situations shall staid out clearly from the rest, and how these
salient points shall be grouped and linked by the connecting
threads of association and shall coalesce into effective wholes.
And if in the comparatively helpless human infant the
congenital modes of response seem less organized than those
of the chick, if there is a larger percentage of random and
apparently aimless movements, if the organic management of
the bodily estate is less definitely ordered by the terms of the
hereditary bequest, if there is more of maternal guidance and
fosterage ; still the data are provided in a substantially similar
way. The situations are indeed destined to become more
complex, the distinctions which arise in consciousness are more
numerous, the coalescence and association include a wider range
and succession of salient points ; a longer time is required to
become acquainted with the transactions of a business con-
ducted in a far greater number of centres: but, at least in the
early stages, the data are of the same kind, and are emphasized
in the same way. Presentation and re-presentation play a

EARLY STAGES OF MENTAL DEVELOPMENT 55

similar ré/e; and the chief difference lies in the fact that
less stereotyped congenital behaviour is supplemented by
some guidance, probably far less than is generally supposed,
from those who lovingly minister to the course of infant
development.

No attempt can here be made to trace even in outline (an
outline which must in any case be imaginary and conjectural)
the sequence of situations which marks the course of mental
development in its earlier stages. An example may, however,
serve to show how the exercise of congenital tendencies may
give rise to a new situation, and lead to a further development
of behaviour.

Ikept some young chicks in my study in an improvised pen
floored with newspaper, the edges of which were turned up and
supported, to form frail but sufficient retaining walls. One of
the little birds, a week old, stood near the corner of the pen,
pecking vigorously and persistently at something, which proved
to be the number on the page of the turned-up newspaper.
He then transferred his attention and his efforts to the corner
of the paper just within his reach. Seizing this, he pulled at
it, bending the newspaper down, and thus making a breach in
the wall of the pen. Through this he stepped forth into the
wider world of my study. I restored the paper as before,
caught the bird, and replaced him near the scene of his former
efforts. He again pecked at the corner of the paper, pulled it
down, and escaped. I then put him back as far as possible
from the spot. Presently he came round to the same corner,
repeated his previous behaviour, and again made his escape.

Now, here the inherited tendency to peck at small objects
led, through the drawing down of the paper, to a new situa-
tion, of which advantage was taken. The little drama con-
sisted of two scenes, which may be sufficiently described as
“the corner of the pen,” and “the open way,” this being the
sequence in experience. Subsequently the first scene was
again enacted in presentative terms, and there followed first
a re-presentation of scene ii., with its associated behaviour,
and then the presentative repetition of this scene. We may

56 CONSCIOUSNESS

take this as a sample of the nature of a conscious situation
which is effective in guidance. We have seen the nature of
the elements (sensory data, including as essential those sup-
plied by the behaviour itself, with a pleasurable or painful
tone) which enter into such a situation; we have seen that
they owe their primary origin to direct presentation, but that
they may be subsequently introduced indirectly in re-presenta-
tive form ; we have seen that the situation as a whole results
from the coalescence of the data. There only remains the
question how the felt situation takes effect on behaviour.
And to this question, unfortunately, we can give but a meagre
and incomplete reply. All we can say is, that connections
seem to be in some way established between the centres of
conscious control and the centres of congenital response ; and
- that through these channels the responsive behaviour may be
either checked or augmented (as a whole or in part), accord-
ing to the tone, disagreeable or pleasant, that suffuses the
situation. How this is effected we do not fully know.

TI..—Later Puases In Menta, DEVELOPMENT

Some surprise may be felt that in our brief discussion of
the early stages of mental development nothing has been said
of percepts and concepts, nothing of abstraction or generaliza-
tion. The omission is not only due to a desire to avoid the
subtle technicalities of psychological nomenclature. It is
partly due to the wish not to forejudge a difficult question of
interpretation. Spirited passages of arms from time to time
take place between psychologists in opposing camps, as to
whether animals are or are not capable of forming abstract
and general ideas ; and untrained camp followers hang on the
skirts of the fray, making a good deal of noise with blank-
cartridge. The question at issue turns partly on the defini-
tions of technical terms ; partly, when there is agreement on
this point, on the interpretation to be put on certain modes of
behaviour. Nothing seems at first sight much easier than to
say what we mean by an abstract idea or by a general idea.

LATER PHASES IN MENTAL DEVELOPMENT 57

We are thinking about colour, which is both abstract and
general—abstract, because in itself it is a special quality of
visible objects floated off, so to speak, from other qualities,
such as hardness and weight, shape and size; general, because
it includes many different colours in one group. Looking up
at the bookshelves, we see a volume with a red back. We
neglect the shape, the contents, the lettering ; it is the colour
with which we are immediately concerned, which forms an
important feature in the present thought-situation ; and this
is, in virtue of that situation, abstracted from the rest. But
a chick a few days old may have acquired experience of several
kinds of caterpillars much alike in shape and size ; of which, one
kind is ringed with orange and black. And while the others
are eagerly seized, caterpillars of this kind are left untouched.
It is not the size or the shape which is an effective element in
the situation ; it is the peculiar coloration of the cinnabar
caterpillars. Now, does the effectiveness of this quality in
the stimulus justify the inference that the chick forms an
abstract idea of colour? That clearly depends on our defini-
tion of abstract idea, and on our inferences concerning the
nature of the chick’s mind.

A dog lies dozing upon the mat, and hears a step in the
porch without. His behaviour at once shows that this enters
into the conscious situation. There is, moreover, a marked
difference according as the step has the familiar fall of the
master’s tread, the well-known shuffle of the irrepressible
butcher’s lad, or an unfamiliar sound. These several situations
are, without question, nicely distinguished. Let us suppose
the situation of the moment is introduced by a. strange foot-
fall. It seems to suggest man ; but this cannot be any parti-
~ cular man, since he is as yet invisible and is a stranger. Does
the dog, then, frame a general idea of man ? Does the chamois
do so when, bounding across the snow field, he stops suddenly
on scenting the distant footprints of a mountaineer ? Do you
do so when you hear the bleating of an invisible lamb in the
meadow behind yonder wall? Here, again, the answers we
give to these questions depend partly on the exact meaning

58 CONSCIOUSNESS

of the term “ general idea ;” partly on our interpretation of
what passes through the mind of the being concerned. We
have sought, so far, rather to avoid than to answer these
questions. We seem to be on safe ground so long as we
content ourselves with saying that the orange and black of
the cinnabar caterpillar, the strange footfall, or the trail of
the mountaineer, enter as effective elements into the immediate
conscious situation.

But when we pass to the higher phases of mental develop-
ment we can no longer wholly ignore such questions. When
we are dealing with intellectual human beings, there can
be no doubt that they at least are capable of framing, with
definite intention, and of set purpose, both general and abstract
conceptions. And how do they reach these conceptions? By
reviewing a number of past situations, analyzing them, in-
tentionally disentangling and isolating for the purposes of
their thought certain elements which they contain, and
classifying these abstracts under genera and species—that is
to say, into broader and narrower groups. The primary and
proximate object of this process is to reach a scheme of thought
by which the scheme of nature, as given in experience, can be
explained. And, no doubt, underlying this primary object is
the purpose of guiding future behaviour in accordance with
the rational scheme which is thus attained. Man is sometimes
described as pur excellence the being who looks before and
after. All his greatest achievements are due to his powers of
reflection and foresight.

What share the symbolism of speech takes in the process
briefly indicated in the last paragraph is the subject of much
discussion. Without going so far as to urge that the very
beginnings of reflective thought are inexplicable without its aid,
it may be accepted as obviously true that words are a great
assistance. They may be regarded as intellectual pegs upon
which we hang the results of abstraction and generalization.
It may be said that we often think in pictures or images, and
not in words; but the more abstract and general our thought,
the more it is dependent on the symbolic elements.

LATER PHASES IN MENTAL DEVELOPMENT 59

We may say, then, that the higher phases of mental
development are characterized by the fact that the situations
contain the products of reflective thought, presumably absent
in the earlier stages ; they are further characterized by a new
purpose or end of consciousness, namely, to explain the
situations hitherto merely accepted as they are given in pre-
sentation or re-presentation; they require deliberate attention
to the relationships which hold good among the several
elements of successive situations ; and they involve, so far as
behaviour is concerned, the intentional application of an ideal
scheme with the object of rational guidance. We shall follow
Dr. Stout in terming this later stage of mental development
the ideational stage ; and in speaking of the simpler situations
considered in the preceding section as belonging to the
perceptual stage.

It should be observed that we are not attempting to
determine just where, in the scale of organic existence, the line
between the perceptual and the ideational stages of mental
development is to be drawn. We are certainly very far from
asserting that the one does not give rise to the other in the
course of an evolution which is orderly and progressive. We
are merely contrasting the rational guidance of effective
consciousness at its best with the earlier embryonic condition
out of which it has arisen by natural genesis. In doing this
we have been forced to make some reference to the difficulties
of technical nomenclature. And some further reference is
necessary lest our point of view be misunderstood.

We shall regard these abstract and general ideas as the
products of an intentional purpose directed to the special end
of isolating the one and of classifying the other; we shall
reserve the term rational for the conduct which is guided
in accordance with an ideal scheme or deliberate plan of
action ; while for behaviour to the guidance of which no such
reflection and deliberation seems to have contributed we shall
reserve the term intelligent. If, for example, the rejection of
a cinnabar caterpillar by the chick is the direct result of
experience through the re-presentation in the new situation

60 CONSCIOUSNESS

of certain elements introduced during the development of
a like situation, we shall call it an intelligent act. But if we
have grounds for supposing that the situation is reflectively
considered by the chick in relation to an ideal and more or
less definitely conceived plan of action which is (perhaps
dimly) taking form in its mind, we shall regard it as so far
rational. And so, too, in other cases of animal behaviour. Now,
with regard to the control through which consciousness is
effective in the guidance of behaviour, it is necessary, in view
of these considerations, to distinguish its intelligent from its
rational exercise. And this is of importance since we generally
speak of control in the latter sense in reference to human
conduct. Intelligent control (on the perceptual plane) is due
to the direct operation of the results of experience without
the intervention of any generalized conception or ideal. In
rational control (on the ideational plane), such conceptions
and ideals exert a controlling influence. If, to prevent a boy
sucking his thumb we administer bitter aloes, we trust to
intelligent control through the immediate effects of experience ;
but if he be induced to give up the habit because it is babyish,
he so far exercises rational control. What we call self-control
is of this type. Only one more distinction need be drawn.
Intelligent behaviour, founded on direct association gained
through previous experience, we shall attribute to impulse; but
for rational conduct, the outcome of reflection and deliberation,
we seek to ascertain the motive. In human affairs our motives
are referred to certain categories each of which presupposes an
ideal scheme, prudential, esthetic, ethical, or other. To act
from motive and not from impulse is to act deliberately,
because we judge the action to be expedient, seemly, or right,
as the case may be. If, then, we contrast the lower perceptual
stages of mental evolution with the higher ideational phases,
the former includes behaviour due to impulse ; but from it
conduct due to motive is excluded.

‘THE EVOLUTION OF CONSCIOUSNESS 61

IV.—Tue Evouution or Consciousness

The origin of consciousness, like that of matter or energy,
appears to be beyond the pale of scientific discussion. The
appearance of effective consciousness on the scene of life does
indeed seem to justify the belief in the prior existence of sen-
tience as the mere accompaniment of organic behaviour. Ex
nihilo nihil fit. And since effective consciousness must, on this
principle, be developed from something, it is reasonable to
assume that this something is pre-existing sentience. Again,
we may assume that this sentience is a concomitant of il life-
processes, or only of some. But we have no criterion by which
we can hope to determine which of these alternatives is the
more probable.

We appear, however, at all events to have evidence that
when effective consciousness does enter on the scene and play
its part in the guidance of behaviour, its progress is, in tecl-
nical phraseology, marked by that differentiation of conscious
elements, and that integration of these differentiated items,
which are seemingly the correlatives of the differentiation and
integration of nervous systems. There is thus, presumably, a
progressive development of orderly complexity in the conscious
situations of which controlled or guided behaviour is the out-
come. And when this has reached a certain stage—what stage
it is most difficult to determine—the relationships, at first
implicit in the conscious situations, as they naturally arise in
the course of experience, begin to be rendered explicit with the
dawn of reflection. Intentional abstraction and generalization
to which data are afforded by the reiterated emphasis in ex-
perience of the salient features in successive situations, supply
new elements to the more highly developed situations of
rational life. Ideal schemes and plans of action, the products
of reflection and foresight, take form in the mind and enter
into the conscious situation. And the intelligent animal,
hitherto the creature of impulse, guided only by the pleasur-
able or painful tone which gives colour to experience, becomes

62 CONSCIOUSNESS

a rational being, capable of judging how far his own behaviour
and that of others is conformable to an ideal.

If, then, we were asked to characterize in the briefest
possible terms the stages of conscious evolution, we should say
that in the first stage we have consciousness as accompaniment ;
in the second, consciousness as guide ; in the third, conscious-
ness as judge. And if we were pressed to apply distinctive
terms to these three, we should adopt St. George Mivart’s
term consentience for the mid-phase, and speak of mere sentience
in the first stage ; consentience in the second ; and conscious-
ness, with restricted signification, in the third and highest
stage. Such a distinction in ternis is, however, a counsel of
perfection, and we shall not attempt to preserve it in the
following pages, in which the word “ consciousness” will be
used in a comprehensive sense.

CHAPTER III
INSTINCTIVE BEHAVIOUR

I.—DeErinition oF Instinctive BEHAVIOUR

THERE are probably few subjects which have afforded more
material for wonder and pious admiration than the instinctive
endowments of animals. “I look upon instinct,” wrote
Addison in one of his graceful essays, ‘as upon the principle
of gravitation in bodies, which is not to be explained by any
known qualities inherent in the bodies themselves, nor from
any laws of mechanism, but as an immediate impression from
the first Mover and the Divine Energy acting in the creatures.” *
In like manner Spence said : ‘We may call the instincts of
animals those faculties implanted in them by the Creator, by
which, independent of instruction, observation or experience,
and without a knowledge of the end in view, they are all alike im-
pelled to the performance of certain actions tending to the well-
being of the individual and the preservation of the species.” t
According to such views, instinct is an ultimate principle the
natural genesis of which is beyond the pale of explanation.
But similar views were, at the time these passages were written,
held to apply, not only to animal behaviour, but also to animal
structure. The development of the stag’s antler, or of the
insect’s wing, was also regarded as “an immediate impression
from the first Mover and the Divine Energy acting in the
creatures.” This view, however, is, neither in the case of

* Spectator, No. 120.
+ Kirby and Spence, “Introduction to Entomology,’ Letter xxvii.
p. 587 (7th Edit., 1858). ~

63

64 INSTINCTIVE BEHAVIOUR

structure nor in the case of behaviour, that entertained by
modern science. It is indeed an expression of opinion con-
cerning the metaphysics of instinct. Leaving the question of
ultimate origin precisely where it stood in the times of Addison
and of Spence, modern science seeks to trace the natural ante-
cedents of all natural phenomena, and regards structure and
behaviour alike as the products of evolution, endeavouring to
explain the manner of their genetic origin in terms of
progressive heredity.

Omitting, therefore, all reference to problems which, how-
ever important, are beyond the limits of scientific inquiry,*
we may take as a basis for further discussion Spence’s definition,
according to which the instincts of animals are those faculties
by which, independent of instruction, observation, or experi-
ence, and without a knowledge of the end in view, they are all
alike impelled to the performance of certain actions tending to
their own well-being and the preservation of the species.

Let us first consider the reference of instinctive actions to
a faculty by which animals are said to be impelled to their
performance. Paley also defined instinct as “a propensity
prior to experience.” And unquestionably in the popular con-
ception itis usual to attribute instinctive acts to some such
conscious cause. But it will be more convenient, for the
present, to consider instinctive behaviour from the objective
point of view, as it is presented to our observation ; we may then
proceed to the further consideration of the conscious con-
comitants which may be inferred. From the objective point
of view, therefore, we may agree with Professor Groos, who
says t that “the idea of consciousness must be rigidly excluded
from any definition of instinct which is to be of practical
utility,” since “it is always hazardous in scientific investigation
to allow an hypothesis which cannot be tested empirically.”
In this we have the support of Dr. and Mrs. Peckham, whose
studies of the life-histories of spiders and wasps are models of

* Cf. supra, p. 18.
+ “The Play of Animals,” translated by Elizabeth L. Baldwin,
p. 62.

DEFINITION OF INSTINCTIVE BEHAVIOUR 6s

careful and patient investigation. ‘“ Under the term Instinct,”
they say, “we place all complex acts which are performed
previous to experience, and in a similar manner by all members
of the same sex and race, leaving out as non-essential, at this
time, the question of whether they are or are not accompanied
by consciousness.” *

It may be said, however, that some reference to the conscious
aspect of instinctive behaviour is implied by saying that the
acts are performed without instruction or experience. But the
reference at present is wholly negative. We may say, as
the result of observation, that instinctive acts are performed
under such circumstances as exclude the possibility of guidance
in the light of individual experience, and render it in the
highest degree improbable that there exists any idea of the end
to be attained. But this is a very different position from that
of asserting the presence of a positive faculty or propensity
which impels an animal to the performance of certain actions.
This it is which, from the observational point of view, is
unnecessary. For the reference of a given type of observed
behaviour to a “propensity” so to behave or to a “ faculty ”
of thus behaving, is no more helpful than the reference of the
development of any given type of structure toa ‘‘ potentiality ”
so to develop. We may, therefore, without loss of precision,
simplify Spence’s definition by stating that instinctive behaviour
is independent of instruction and experience, and tends to the
well-being of the individual and the preservation of the species.

Let us next consider the clause which affirms that instinctive
behaviour is prior to experience. This is well in line with the
distinction now drawn by biologists between congenital and
acquired characters. It refers them to the former category, and
implies that the organic mechanism by which they are rendered
possible is of germinal origin. This is not, however, universally
admitted. Professor Wundt, for example, approaching the
subject from the point of view afforded by the study of man
and the higher animals, gives to the term a wider meaning,

* George W. and Elizabeth G. Peckham, “On the Instincts and

Habits of the Solitary Wasps,” p. 231.
F

66 INSTINCTIVE BEHAVIOUR

and so defines instinct as to include acquired habits. “ Move-
ments,” he says,* “which originally followed upon simple or
compound voluntary acts, but which have become wholly or
partly mechanized in the course of individual life, or of generic
evolution, we term zstinctive actions.” In accordance with
this definition, instincts fall into two groups. Those “ which,
so far as we can tell, have been developed during the life of the
individual, and in the absence of definite individual influences
might have remained wholly undeveloped, may be called acquired
instincts.” They have become instinctive through repetition.
“To be distinguished from these acquired human instincts are
others which are connate.” Now, there can be no question
that behaviour which has become habitual through frequent
repetition is frequently, in popular speech, described as in-
stinctive. We hear it said that the experienced cyclist guides
his machine instinctively. And the word is similarly used in
many like cases. But we shall find it conducive to precision
and clearness of thought to emphasize the distinction between
what is acquired in the course of life and what is congenital in
the race. And to this end we shall regard behaviour which
has ‘become mechanized in the course of individual life” as
due to acquired habit, reserving the term instinctive for such
behaviour as is independent of individual experience. We
shall, in short, so far accept Spence’s definition.

In this definition, as in those of the majority of naturalists,
it seems to be further implied that instinctive behaviour is of
a relatively definite kind, though it is no doubt subject to such
variation as is found in animal structure and organization.
Mr. Rutgers Marshall, however, in a recent work,{ protests
against any such implication, and urges that “this variableness
is so wide that definiteness of reaction cannot for a moment be
used as a differentia in relation to instinct without narrowing
our conception of the bounds of instinct in a manner to be
deplored.” “The actions,” he says, “connected with the
preparation for self-defence, those connected with protection of

* « Lectures on Human and Animal Psychology,” pp. 388, 397, 399.
+ “Instinct and Reason,” pp. 90, 92.

w

DEFINITION OF INSTINCTIVE BEHAVIOUR 67

the young, with nest-building, with migration, etc., these
actions are surely to be classed as instinctive ; and yet they are
exceedingly variable and unpredictable in detail ; all that we
can predict is the general trend of the varying actions which
result from varying stimuli under varying conditions, and
which function to some determinate biological end.”

Mr. Marshall then proceeds to argue that we are “ warranted
in speaking of the ethical instincts, of the patriotic instincts,
of the benevolent instincts, and of the artistic instincts ;” and
thus leads up to the position, to be further elaborated in his
work, that there exists in man a religious instinct which has
fulfilled a function of biological value in the development of
our race. Now, here again there is much in popular usage of
the words instinct and instinctive which lends support, for
what it is worth, to Mr. Marshall's very broad conception of
the range of instinct. Again and again we hear, in the pulpit
and elsewhere, of the religious instinct ; we hear, too, of the
benevolent, patriotic, and artistic instincts, and more besides.
But what we are endeavouring to define is a type of behaviour
which, as such, is prior to instruction and experience. Can we
affirm that patriotic and religious behaviour conforms to such
a type? Is it unquestionably congenital and not acquired ?
If we are forced to give negative answers to these questions we
must regard Mr. Marshall’s conception of instinct (one inclusive
of multifarious tendencies which have a biological value) as
too broad and too vague to be of any service to us at this stage
of our study of animal behaviour.

What, then, shall we understand by Spence’s phrase that
instinct involves the performance of “certain actions” ? And
how far shall we accept it ? We shall take it as implying so
much definiteness of behaviour as renders instinctive acts sus-
ceptible of scientific investigation, and in this sense shall
accept it with some modification of phraseology. We shall
freely admit, however, the existence of variations of instinctive
behaviour analogous to variations in animal structure. It is
the occurrence of such variations that renders the natural

selection of instinctive modes of behaviour conceivable. We

68 INSTINCTIVE BEHAVIOUR

shall also admit some, nay much, variation in detail. Take, for
example, two of the cases which Mr. Marshall cites—nest-
building and migration. Both involve, not merely a simple
response to a given stimulus, but a complex sequence of actions.
In detail there may be much variation even among members of
the same species. And yet, can it be questioned that the
behaviour as a whole is in each case relatively definite ? May
we not even say that it is remarkably definite? May we not
even go further, and assert that only on the assumption that
instinctive behaviour is relatively definite, can we regard it
as a subject for scientific investigation, and can we hope to
distinguish it from other modes of behaviour ?

The next point for consideration in Spence’s definition,
which we have taken as our text, is his characterization of
instinctive acts as ‘“‘ tending to the well-being of the individual
and the preservation of the species.” Here we have Mr.
Marshall with us, for he too lays stress on the fact that
instinctive behaviour has reference to a definite biological end.
But in saying that the biological end is the objective mark of
an instinct,* he seems to bein error. Because, in the first place,
there are other “ objective marks,” and because, in the second
place, this objective mark is not restricted to instinctive
behaviour. According to Spence, a further characteristic of
instinctive acts is that they are independent of instruction or
experience ; and this serves to differentiate them from other
modes of behaviour which are also subservient to a biological
end. Intelligent behaviour, not less than that which we term
instinctive, has reference to a biological end. Many intelligent
acts, for example, have for their object the well-being of the
individual ; many subserve race preservation ; these bear, every
whit as much as instinctive acts, the ‘‘ objective mark” which
Mr. Marshall regards as characteristic of instinct. And if we
turn to his subjective criterion—the absence of any conception
of the biological end which the behaviour subserves—Mr.
Marshall’s position is equally untenable. There are thousands
of acquired modes of behaviour, dependent on instruction or

* « Instinct and Reason,” p. 91.

DEFINITION OF INSTINCTIVE BEHAVIOUR 69

experience, in which there is, on the subjective side, so far as
we can judge, no conception of the biological end to be attained.
What can the animal in the early stages of intelligence know
of biological ends? Mr. Marshall’s subjective criterion applies
just as much to a wide range of intelligent behaviour as it does
to instinctive actions.

In accepting, therefore, Spence’s statement that when animals
behave instinctively they perform, without a knowledge of the
end in view, certain actions tending to their own well-being
and the preservation of the species, we must take it in con-
nection with the preceding limitation, remembering that they
are also performed without instruction and experience.

A further point for very brief consideration is suggested by
the phrase in which Spence says that animals are all alike im-
pelled to the performance of certain actions. As it stands it
is too sweeping and general. Still, we do require some explicit
statement of the facts which he had in mind when he wrote
the words “all alike.” And we find it with sufficient exactness
in Dr. Peckham’s definition, where he comprises under the
category of instinctive behaviour “ all complex acts which are
performed previous to experience, and i @ similar manner by
all members of the same sex and race.” This places congenital
behaviour in line with morphological structure as a subject for
comparative treatment.

One more question remains. What shall we understand
by ‘complex acts” ? In the first place, it is well to restrict
the term instinctive to co-ordinated actions ; and this implies
the presence of nerve-centres by which the co-ordination is
effected. We thns exclude.the organic behaviour of plants,
since there is no evidence in the vegetable kingdom of
co-ordinating centres. In the second place, the co-ordination
is, as we have seen, congenital, and not acquired in the course
of individual experience. Young water-birds, and indeed
young chicks, as soon as they are born, and have recovered
from the shock of birth, can swim with definite co-ordination
of lee movements. Here the definiteness is not only congenital,
but connate, if we use the latter term for an instinctive activity

70 INSTINCTIVE BEHAVIOUR

which is performed at or very shortly after birth. On the
other hand, young swallows cannot fly at birth ; they are then
too immature, and their wings are not sufficiently developed.
But when they are some three weeks old, and the wings have
attained functional size and power, little swallows can fly with
considerable if not perfect skill. The co-ordination is con-
genital, for it is not acquired in the course of individual
experience ; but it is not connate, since it is not exhibited at
or shortly after birth. The term deferred may be applied to
such congenital activities as are thus carried out when the
animal has undergone a certain amount of further develop-
ment after birth.

In the third place, it is customary to distinguish between
such reflex actions as have already been briefly exemplified,*
and instinctive behaviour. It is, however, by no means easy,
if indeed it be possible, to draw any sharp and decisive line of
demarcation. Instinct has indeed been well described by Mr.
Herbert Spencer as compound reflex action ; hence the dis-
tinction between instinctive. and reflex behaviour turns in
large degree on their relative complexity. It would seem,
however, that whereas a reflex act—such as the withdrawal of
the foot of a sleeping child when the sole is tickled—is a
restricted and localized response, involving a particular organ
or a definite group of muscles, and is initiated by a more or
less specialized external stimulus ; instinctive behaviour is a
response of the animal as a whole, and involves the co-opera-
tion of several organs and of many groups of muscles. Partly
initiated by an external stimulus or group of stimuli, it is also,
seemingly, determined in part, in a greater degree than reflex
action, by internal factors which cause uneasiness or distress,
more or less marked, if they do not find their normal in-
stinctive satisfaction. This point, however, may be more
profitably discussed in connection with the conscious aspect of
instinct. If, then, we say that reflex acts are local responses of
the congenital type due to specialized stimuli, while instinctive
activities are matters of more general behaviour, usually

* Chapter I., Section Y.

INSTINCTIVE BEHAVIOUR IN INSECTS 71

involving a larger measure of central (as opposed to local or
ganglionic) co-ordination, and due to the more widely-spread
effects of stimuli in which both external and internal factors
co-operate, we shall probably get as near as is possible to the
distinction of which we are in search. But it must be remem-
bered that there are cases in which the distinction can hardly
be maintained.

We are now in a position to define instinctive behaviour
as comprising those complex groups of co-ordinated acts which
are, on their first occurrence, independent of experience ;
which tend to the well-being of the individual and the pre-
servation of the race; which are due to the co-operation of
external and internal stimuli ; which are similarly performed
by all the members of the same more or less restricted group
of animals; but which are subject to variation, and to sub-
sequent modification under the guidance of experience.

II.—InstinctivE BEHAviouR In INSECTS

Since instinctive behaviour is, by definition, independent
of experience, and since the animals which act instinctively
are also, in many cases, able to act intelligently, it is clear
that, apart from hereditary variations, we must expect to find
acquired modifications of instinct. As Huber said of bees,
their instinctive procedure often indicates “a little dose of
judgment.” It is, indeed, exceedingly difficult, as a matter of
observation, to distinguish between hereditary variation and
acquired modification. For the réle played by these two
factors in any given behaviour can only be determined if the
whole life-history of the individual be known, and if thére be
opportunities for comparing it with the complete life-histories
' of other members of its race. And this is seldom possible.
These considerations must be borne in mind as we proceed
to a brief study of some of the instinctive modes of behaviour
in insects.
Dr. and Mrs, Peckham’s investigations on the instincts and

“72 INSTINCTIVE BEHAVIOUR

habits of the solitary wasps have been described in a volume *
worthy to be placed by the side of Fabre’s “Souvenirs.” Their
descriptions seem to glow with the warm sunshine, and are
redolent of the fresh air which afforded the conditions under
which the observations were conducted. We can but regret
that, in extracting from their bright pages some of the salient
facts, the natural delicacy and grace of their treatment must
be lost. For we can only give the dry skeleton which they
have clothed with the flesh of lively detail. They enumerate
the following primary modes of instinctive behaviour :—

1. Stinging.

2. Taking a particular kind of food.

3. Method of attacking and capturing prey.

4. Method of carrying prey.

5. Preparing nest, and then capturing prey, or the
reverse. ‘
6. The mode of taking prey into the nest.

7. The general style and locality of the nest.

8. The spinning or not spinning of a cocoon, and its
specific form when one is made.

When the young Pelopeus emerges from the pupa-case

and gnaws its way out of the mud cell, with limp and flaccid
wings, it responds to a touch by well-directed movements of
the abdomen with thrusts of the sting, as perfect as those of
the adult. There is clearly no opportunity here for either
instruction or experience to afford any intelligent guidance.
Stinging is an instinctive act. And it is an act of which great
use is made in the capture of prey which shall serve for food
to the young—it has a biological end. But the wasps of
different species do not have to learn by experience what prey
to attack. It is by instinct, too, that they take their proper
food-supply, one caterpillars, another spiders, a third flies or
beetles. So deeply seated, indeed, is the hereditary preference,
that no fly-robber ever takes spiders, nor will the capturer of
spiders change to caterpillars or beetles. Some keep to a few

* “On the Habits and Instincts of the Solitary Wasps,” by George W.
and Elizabeth G. Peckham (1898),

INSTINCTIVE BEHAVIOUR IN INSECTS 73

species or genera, while Philanthus punctatus preys chiefly or
entirely on bees of the genus Hualictus.

Romanes * thought that the manner of stinging and para-
lyzing their prey might “be justly deemed the most remarkable
instinct in the world.” Spiders, insects, and caterpillars are
stung, he says, ‘in their chief nerve-centres, in consequence
of which the victims are not killed outright, but rendered
motionless ; they are then conveyed to a burrow and, continu-
ing to live in their paralyzed condition for several weeks, are
then available as food for the larvee when these are hatched.
Of course the extraordinary fact which stands to be explained
is that of the precise anatomical, not to say also physiological
knowledge which appears to be displayed by the insect in
stinging only the nerve-centres of its prey.” Eimer + thought
that it “is absolutely impossible that the animal has arrived
at its habit otherwise than by reflection upon the facts of
experience.” “At the beginning,” he says, “she probably
killed larvee by stinging them anywhere, and then placed them
in the cell. The bad results of this showed themselves ; the
larvee putrified before they could serve as food for the larval
wasps. In the mean time the mother wasp discovered that
those larvee which she had stung in particular parts of the
body were motionless but still alive, and then she concluded
that larve stung in this particular way could be kept for a
longer time unchanged as living motionless food.”

Now, since these wasps, when they have stored their nests
and laid an egg on one of the victims, close it up once and for
all, and take no further interest in it or its contents, there
seems no opportunity, at any rate in the existing state of
matters, for the acquisition of that experience on which Eimer
relied. But both his explanation and Romanes’s difficulty are
based on the following assumptions: first, that the victims
are instinctively or habitually stung in the chief nerve-centres ;
secondly, that when thus stung they are not killed but remain
paralyzed for weeks ; and thirdly, that the marvellously definite

* “Mental Evolution in Animals,” p. 299.
+ “Organic Evolution,” translated by J. T. Cunningham, p. 280.

74 INSTINCTIVE BEHAVIOUR

and delicate instinctive behaviour is in direct relation to the
uniform result of prolonged paralysis and consequent preserva-
tion of the food in the fresh state. But Dr. Peckham’s careful
observations and experiments show that, with the American
wasps, the victims stored in the nests are quite as often dead
as alive ; that those which are only paralyzed live for a vary-
ing number of days, some more, some less ; that wasp larvee
thrive just as well on dead victims, sometimes dried-up, some-
times undergoing decomposition, as on living and paralyzed
prey ; that the nerve-centres are not stung with the supposed
uniformity ; and that in some cases paralysis, in others death,
follows when the victims are stung in parts far removed from
any nerve-centre. ‘‘ We believe,” he says, ‘that the primary
purpose of the stinging is to overcome resistance, and to pre-
vent the escape of the victims, and that incidentally some of
them are killed and others are paralyzed.”

If, therefore, as will probably be shown to be the case,
these conclusions are found to be generally true for this in-
teresting group of insects, the mystery of “the precise ana-
tomical, not to say also physiological knowledge which appears
to be displayed” by these wasps turns out to be one of our
own fabrication. It melts away in the light of fuller and
more searching investigation.

It must not be supposed, however, from what has been
said, that the behaviour in the act of stinging is altogether
indefinite. On the contrary, each species proceeds in a rela-
tively definite manner with some variation or modification of
method. Philanthus punctatus, for example, stings the bees,
on which she preys, under the neck, and the thrust is at once
fatal. Dr. Peckham further notes that he was only suc-
cessful in getting the wasps to sting when they were hunting ;
those that had not yet begun to store the nests paid no
attention to the bees. This is an example of that internal
factor to which reference was made in the last section.
Marchal observed that Cerceris ornata runs the end of her
abdomen along the under surface of the thorax of the bee, and
delivers her thrust at the division of the segments—that is,

INSTINCTIVE BEHAVIOUR IN INSECTS 75

where the sting can enter. The action does not imply any
physiological knowledge. In general she begins at the neck.
Spiders are usually, but not always, stung on the ventral
surface. To give but one more example, Dr. Peckham ob-
served in three cases the procedure of Ammophila urnaria
which preys on caterpillars, and often, after stinging, bites the
neck in several places, this process being termed malaxation.
In three observed captures, all the caterpillars being of the
same species and alike in size, the thrusts were given on the
ventral surface near the middle line, between the segments,
In the first, seven stings were given at the extremities (there
being thirteen segments), the middle Segments being left
untouched, and no malaxation was practised. In the second,

Fic. 11.—Solitary Wasp stinging Caterpillar (after Peckham),

seven stings were again given, but in the anterior and middle
segments, followed by slight malaxation. In both these cases
the first three thrusts were in definite order, behind the third,
the second, and the first segments successively. In the case
of the third caterpillar, only one thrust was given, between the
third and fourth segments—that ‘is to say, in the position of
the first stab in the other cases,—and after this one thrust
there was prolonged malaxation. Of fifteen stored caterpillars
examined, some lived only three days, others a little longer,
while a few showed signs of life at the end of a fortnight. In
more than one instance the second of the two caterpillars

76 INSTINCTIVE BEHAVIOUR

stored in each nest died and became discoloured before the
first one was entirely eaten. The larva under such circum-
stances ate it with good appetite, and then spun its cocoon as
if nothing unpleasant had occurred.

The mode of carrying their booty is in these wasps in-
stinctive, and relatively uniform. Ammophila urnaria grasps
the caterpillar, near the anterior end, in her mandibles, and
carries or drags it beneath her legs, walking forwards. It is

Fic. 12.—Solitary Wasp dragging a Caterpillar to its Nest (after
Peckham).

generally but not always with the ventral surface uppermost.
Pompilus takes hold of her spider anywhere, but always drags
it over the ground, walking backwards. Ozybelus clasps her
fly with her hind legs; ember with the second pair. Each
works after her own fashion in a way that is relatively uniform
for each species.

The general style of the nest, its mode of construction, and
its method of closure, are always performed, says Dr. Peck-
ham, by each species in a similar manner, not indeed in
circumstantial detail, but quite in the same way in a broad
sense. Variation or modification is always present, but the
tendency to depart from a nest of a given type is not ex-
cessive. Some dig in the ground curved tunnels, with or

INSTINCTIVE BEHAVIOUR IN INSECTS 77

without one or more chambers. Others bore into decaying
wood ; others use straws, or make tunnels in bramble stems ;
while the mud-daubers build cells in which to store the
food and lay the egg. This is sometimes deposited on the
first, sometimes on the last, sometimes on some intermediate
victim, but generally in much the same place and position.
Ammophila, for instance, lays it on the side of the sixth or
seventh segment—that is to say, in about the mid position.

Some species first capture their prey, and then make the
nest in which it is to be entombed. Others first prepare the
nest, and then carry or drag their prey to it—often from con-
siderable distances—quite irrespective of what seems to us
the more appropriate method of the two under the particular
circumstances of the case. And the way in which the victim
is dragged into the nest is similarly a matter of inheritance.
Each way is characteristic of the species concerned, and would
be an important part of any definition of the animal based
upon its modes of behaviour. For example, a Sphex places
her grasshopper just at the entrance of the nest, which she then
enters herself before dragging in her prey by the antenne.
When the wasp was in the hole, Fabre moved the victim a
little way off ; the wasp came out, brought the grasshopper to
the entrance as before, and went in a second time. This was
repeated about forty times, each time with the same result,
until the patience of the naturalist was exhausted, and the
persistent wasp took her booty in after her appropriate fashion.
She must place the grasshopper close to the opening; she
must then descend and examine the nest, and, after that, must
drag it down. Nothing less than the performance of these
acts in a certain order satisfies her instinctive impulse.

In a private letter, from which he kindly allows me to
quote, Dr. Peckham says: “We have recently made some
experiments on this wasp (Sphex ichneumonea). First we
allow her to carry in her prey undisturbed, to see how far she
was faithful to the traditions of her ancestors, and to observe
ber normal methods. On the next day, when she had placed
her grasshopper just at the opening of the nest, and while she

78 INSTINCTIVE BEHAVIOUR

was below, we drew it back to a little distance. She came out,
and we both repeated our operations four times—she running
down into the nest, always after getting the grasshopper into
position, and we as regularly drawing it away. The fifth
time she changed her plan, seized it by the head and backed
into the nest with it. The next day, at the fourth trial, she
straddled it and walked head first into the nest with it; and
on the fourth day, at the eighth trial, she backed in with it as
on the second day.” These interesting observations show
that the wasp has sufficient intelligence to modify her pro-
cedure in accordance with an unwonted situation. The
_ “consecutive necessity,” as it has been termed, has a potent
influence, but is not absolute.

Fabre notes a case of similar consecutive necessity in the
case of the mason bee, Chalicodoma. If while a bee is pro-
visioning its nest with honey and pollen the structure be
destroyed, she sometimes breaks open a completed cell, and,
having done so, goes on bringing more provision, though the
cell already contains a sufficient store of food ; and only when
she has completed the superfluous storing does she deposit her
egg and seal up the cell. So, too, when the cell is removed
in an early stage of construction, and another completed cell
already partially stored is substituted, the bee, instead of simply
adopting the new cell, goes on building until the cell is as
much as one-third beyond the usual height ; then, and not till
then, does she proceed in due course to the next stage of the
instinctive procedure, the provisioning of the cell.

From our general knowledge of animal nature, we should
expect to find parasitic forms ready to take advantage of the
material stored by such insects as the solitary wasps and the
mason bees. It is said that Chalicodoma provides nourishment
to the larvee of some sixteen unbidden guests. <A parasitic
bee (Stelis nasuta) breaks open a closed cell, and, after deposit-
ing its eggs, seals it up again with mortar. Since her eggs
and larvee develop more rapidly than those of the mason bee,
they are first served with the store of provision, while the
rightful owner is done out of its inheritance. By a curious

INSTINCTIVE BEHAVIOUR IN INSECTS 79

act, of what appears to us like retributive justice, these para-
sitic larves sometimes fall a prey to another parasite, also a
hymenopterous insect named Jonodontomerus, the larve of
which prey on the young of both bees. Another genus of the
same family, Lewcopsis (Fig. 13, F), also succeeds in piercing with
its ovipositor, at a suitable spot, the walls of the Chalicodoma
cell, and suspends its curious hooked egg (Fig. 13, @) on the
delicate cocoon within which the chrysalis lies. Fabre found
in some cases as many as five of these parasitic eggs on a single
cocoon. But he never found more than one larva in any cell
that he examined. The following is an epitome of his con-
clusions and inferences. From the parasitic egg is hatched a
minute arched grub, with relatively large head and mandibles,
and provided with a number of bristles, which aid it in pro-
gression (Fig. 13, H). It does not, however, at once attack the
bee larva, but makes a series of excursions, the object of which
is to reach und destroy any other parasitic eggs. This was
not actually observed, but the eggs were found to have been
destroyed, and there was seemingly no other means of destruc-
tion under the conditions maintained. The larva, this done,
changes its skin and takes on a new form, destitute of bristles,
with a very small head and minute mandibles (Fig. 13, 1).
In this new form it attacks the Chalicodoma larva, making
avery small incision, through which the juices of the host
are transferred to the guest without further injury to the
grub. It is interesting to note that, if the facts are accurately
described and the inferences are correct, there are associated
with two types of instinctive behaviour two distinct types
of structure. The creature can have no conscious control
over its structural development, and there is no ground for
assuming that it has any control over its instinctive behaviour.

The specialization of structure and of instinctive behaviour,
in accordance with a definite sequence of life-conditions, is
even more remarkable in another of the many parasites which
Chalicodoma unwittingly labours to nourish. This time it is
a fly (Argyromeba), which lays a minute egg on the outside
of the cell. From this egg is hatched a slender threadlike

Fic. 13.—Insect Larve. a, B, of Sitaris ; c, D, B, of Argyromeba ; G, H, 1,
of Leucopsis ; ¥, imago of Leucopsis (after Fabre).

INSTINCTIVE BEHAVIOUR IN INSECTS 81

worm, barely one-twentieth of an inch in length (Fig. 13, Cc).
It has three pairs of longish bristles near the anterior end, and
a single yet longer pair at the hinder extremity. These aid
it in creeping over the wall of the cell. Its small head is
armed with short, stiff bristles. For many days it wanders
over the surface of the cell, inserting its bristly head into each
minute cranny and crack. Throughout this long period it
has never a bite nor sup. Probably many of them never
succeed in finding a crevice by which they can effect an
entrance, but those that do manage to wriggle in undergo a
change, lose their bristles, and develop a minute suctorial
mouth, through which the contents of the larva are absorbed
into their swelling bodies (Fig. 13, p). When fully grown
they are quite helpless, and unable to get out from the cell in
which they arenow imprisoned. For months they lie quiescent,
but in the succeeding spring they pass into a pupal condition
_ very different from that of most flies. The relatively large
head is armed with strong spines; the middle region bears
bristles directed backwards ; the posterior end has short spines
(Fig. 18, E). Fixing itself to the interior of the cell by the
latter, it strikes with its armoured head repeated blows on
the walls of its prison until a breach is at last made, and
sufficiently enlarged to form a suitable exit. Then the pupa-
skin bursts, and the imago insect emerges and flies off. At
each stage of life there is the closest relation between structure
and behaviour, and each is equally adapted to a biological end
of which the creature has never had an opportunity of gaining
any experience.

Exceedingly multifarious are the ways in which insects
thus provide for the future of young they will never see.
Antherophagus lives in flowers, and is believed to seize with its
mandibles humble bees, which then unwittingly bear the
parasitic beetle to the nests in which alone the larve have
been found. The larve of our common oil-beetle (Jeloé) are
parasitic on the bee, Anthophora. It deposits its ten thousand
eggs without observable discrimination ; but the active young
larva instinctively seizes and attaches itself to any hairy object.

@

82 INSTINCTIVE BEHAVIOUR

Thousands must go astray. They have been found on hairy
beetles, flies, and bees of the wrong genus. Some, however,
become thus attached to the one suitable species, and are con-
veyed by the Anthophora to her nest, where they promptly eat
the egg she lays. It is not difficult to picture to one’s self how
this incompletely evolved instinct might be further perfected
by natural selection, through the survival of those females
which laid their eggs in the haunts of the bee-host. And
such an advance in instinctive behaviour is seen in another
and rarer beetle—Situris. Her eggs are laid in August near
the entrance to a nest of the Anthophora. In September they
hatch to form larvee, which hibernate in groups till the following
spring. Then they become active (Fig. 13, a), and attach
themselves to hairy objects. Being near the Anthophora nest,
there is an increased chance of their fastening upon this bee.
The chance is still far from good, for if this were so, we should
not find that the Sidaris laid as many as two thousand eggs.
Still, on these grounds, we may presume that its chance of
survival ig about five times as good as that of Meloé, which
lays ten thousand eggs. The larva is said generally to attach
itself to a male bee, which is hatched earlier than his mate,
and to pass on to the female at the nuptial period ; but in any
case it eventually slips on to the egg that she lays. This
forms the food of the larva during the remainder of this stage
of its existence. It then moults and assumes a new form,
capable of feeding on the honey (Fig. 13, B) ; and, after further
changes, becomes a pupa, and then assumes the imago condition.

In these cases the advantage is wholly on the side of the
parasite. But there are cases of close relationship between
insects and flowering plants where the instinctive behaviour
gives rise to reciprocal benefit. The Yucca is a genus of
American Liliaceous plants, with large pale sweet-smelling
flowers; and these are dependent for fertilization on the
instinctive behaviour of a small straw-coloured moth of the
genus Pronuba. Just when the Yucca plant blossoms in
the summer, the moths emerge from their chrysalis cases.
They mate; and the female then flies to a flower, collects a

INSTINCTIVE BEHAVIOUR IN INSECTS 83

pellet of pollen from the anthers, proceeds to another flower,
pierces the pistil with her sharp ovipositor, lays her eggs
among the ovules, and finally darting to the stigma stuffs the
pollen pellet into its funnel-shaped extremity (Fig. 14). If
the flower be not thus fertilized
the ovules do not develop; and
if the ovules do not develop the
grubs which are hatched from the
moth’s eggs die of starvation.
There are enough ovules to supply
food to the grubs, and leave a
balance to continue the race of
Yuccas.

Whether the female moth is
attracted to the flower by sight
or smell, we do not know. And _ Fic. 14.—Yucca Flower and
whether the male finds the female, ets
in the case of the Yucca moth, through scent, we are not
in a position to state with certainty. It has, however, been
shown that in certain moths* some odour emitted by the
female is the attractive stimulus, affecting sense-organs situated
onthe antenne of the male. To females confined in an opaque
vessel over the mouth of which gauze was tied, the males came
in numbers; but when a clear glass vessel was inverted, and
sand was packed round the mouth, so as to prevent the escape
of air from the interior, no males came, though the imprisoned
females were clearly visible. If the antenne of the males were
either removed or coated with shellac the moths failed to
notice the females even when close to them. In what way the
intact male is made aware of the direction from which the
scent comes, we do not know—possibly by differential stimula-
tion in the antenne, the moth instinctively turning in the
direction of greater stimulation. It will be seen, therefore,
that in the case of the behaviour of the Yucca moth—
behaviour which is essential to the biological end of repro-

* See A. G. Mayer “On the Mating Instinct of Moths.” Ann, and
Mag. of Nat. Hist,, ser. 7, vol. v., Feb., 1900, p. 183,

84 INSTINCTIVE BEHAVIOUR

duction—there is much detail concerning which we are
ignorant. But for our present purpose the important point to
notice is that the procedure of the female cannot be due to
imitation ; nor can it be the outcome of individually acquired
experience ; for the method of procedure is not gradually
learnt, but is carried out without apparent hesitation the first
and only time the appropriate occasion presents itself. Not
only does the moth take no heed of her grubs, but they are so
placed that she could not in any case ascertain by observation
that only if the ovules are fertilized do her offspring thrive.
She cannot possibly know what effect the stuffing of the pollen
on to the stigma exercises, or indeed whether it have any effect
at all. And yet generation after generation these moths collect
the pollen from the anthers and bear it to the stigma. Spence’s
words “without knowledge of the end in view” are amply
justified in this case, as in other cases of typically. instinctive
behaviour.

JII.—Tue Instinctive BrHAviour oF Youne Brrps

Since it is easy to hatch birds of many species in an
incubator, and to rear them under conditions which not only
afford facilities for observation but exclude parental in-
fluence, their study has special advantages. One can with
some approach to accuracy distinguish the instinctive from
the acquired factors in their behaviour.*

The callow young of such birds as pigeons, jays, and
thrushes are hatched in a helpless condition, and require
constant and assiduous ministration to their elementary
organic needs. Most of their instincts are of the deferred
type. But pheasants, plovers, moor-hens, domestic chicks,
and ducklings, with many others, are active soon after birth,
and exhibit powers of complex co-ordination, with little or
no practice of the necessary limb-movements. They walk

* Some of the observations on which the summary of results given in

this section are founded are presented in some detail in “Habit and
Instinct,” pp. 29-100.

INSTINCTIVE BEHAVIOUR OF YOUNG BIRDS 85

and balance the body so soon and so well as to show us that
this mode of procedure is congenital, and has not to be
gradually acquired through the guidance of experience.
Young water-birds swim with neat and orderly strokes the
very first time they are gently placed in water. Even little
chicks a day or two old can swim well. Dr. Thorndike, who
draws attention to this fact,” appears to accept the view,

Fic. 15.—Newly-hatched Chick swimming.

suggested by Dr. Bashford Dean, that the movements are not
those of swimming but only of running. I have carefully
watched the action through the glass walls of a tank and
compared it with that of a young moor-hen. In the two cases
it is quite similar in type, and the type appears to be different
from that of running, though it is perhaps hard to distinguish

* Psychological Review, May, 1899, p. 286.

86 INSTINCTIVE BEHAVIOUR

the two. In any case, the hand over hand action is well
co-ordinated, and is very different from a mere excited struggle.
Chicks twenty-six hours old taken straight from the incubator
drawer, before they had taken food, made directly for the side
of the tank and tried to scramble out. They gradually sank
deeper through the wetting of the down, but could keep afloat
for from two to three minutes. I have made observations on
chicks of various ages from twenty-four hours to a month, and
find in all cases similar results ; but with the older birds the
flapping of the wings and more vigorous action cause them to
get water-logged more rapidly. There issome apparent distress
with cries ; but less than one might expect under the circum-
stances. For the purposes of the above illustration Mr.
Charles Whymper had before him a sketch I made of the
leg-action, and instantaneous photographs of the chicks
swimming for which I am indebted to my colleague Mr.
George Brebner. I have not observed the behaviour of an
adult hen when placed in the water. Dr. Thorndike says,
“there is no vigorous instinct to strike out toward the
shore,” she “will float about aimlessly for awhile and only
very slowly reach the shore.” But Mrs. Foster Wood informs
me that she has seen a hen leap into a pond after her brood of
ducklings and swim to the other side, a distance of twenty
feet.

Diving, in water-birds, is also an instinctive mode of
behaviour; and this is obviously a more difficult procedure
than swimming, one further removed from reflex action.
And careful observations have placed beyond question the fact
that flight is also instinctive. A swallow, for example, taken
from the nest under conditions which made it practically
certain that it had never yet taken wing, exhibited guided
flight, and attempted to alight on a suitable ledge. Of course
flight is generally a deferred instinct, and is not performed
until the wings have reached a suitable state of development.
An instinctive response, which may perhaps be regarded as
one of its initial stages, is seen in quite young chicks. If
placed in a basket, and rapidly lowered therein through a foot

INSTINCTIVE BEHAVIOUR OF YOUNG BIRDS 87

or two, the chick will extend its skinny and scarcely feathered
wings. But though, from the usual conditions of development,
flight in birds is a deferred instinct, yet in exceptional cases
it may be connate. The mound-builders (Megapodes) of the
Australian region are hatched from large eggs in warm earth

Var
Yo Gn
Gee

Fic. 16.—Nestling Megapode, to show the well-developed wings. (From
Dr. R. Bowdler Sharpe’s “ Wonders of the Bird World.”’)

or sand, and are not tended by the parents. So well fledged
are these birds that they can fly the day they emerge from the
egg. Dr. Worcester, while digging in one of their mounds,
made an unsuccessful attempt to seize one which was newly
hatched ; but it flew several rods into thick brush, and this

838 INSTINCTIVE BEHAVIOUR

notwithstanding the fact that it had probably never before
seen the light of day.

It must not be supposed that, in adducing flight as an
example of instinctive behaviour in birds, we are contending
that it is this and nothing more throughout life. The inference
to be drawn from the facts of observation is rather that
instinct provides a general ground plan of behaviour which
intelligent acquisition, by enforcing here and checking there,
perfects and guides to finer issues. Few would contend that
the consummate skill evinced in fully developed flight at its
best, the hurtling swoop of the falcon, the hovering of the
kestrel, the wheeling of swifts in the summer air, the rapid
dart and sudden poise of the humming bird, the easy sweep
of the sea-gull, the downward glide of the stork—that these
are, in all their exquisite perfection, instinctive. A rough but
sufficient outline of action is hereditary; but the manifold
graces and delicacies of perfected flight are due to intelligent
skill begotten of practice and experience.

There are many little idiosyncracies and special traits
of flight which are probably instinctive—such as enable an
ornithologist or a sportsman to recognize a flying bird from
a distance. And the same is true of other modes of behaviour.
The observer of young birds cannot fail to note and to be
impressed by many of these. The way in which a little moor-
hen uses its wings in scrambling up any rough surface is very
characteristic ; so, too, is the manner in which a guinea-chick
runs backwards and then sideways at a right angle when
one attempts to catch him. If suddenly startled, moor-hens
and chicks scatter and hide; plovers drop and crouch with
their chins on the ground; pheasants stand motionless and
silent. Knowledge of the ways of birds enables one to predict
with tolerable accuracy how each kind will behave under given
circumstances. That the actions are always precisely alike
cannot be said with truth; but that the behaviour is so
relatively definite as to be readily recognizable can be con-
fidently asserted. That a moor-hen will flick its tail, that
a chick will dust itself in the sand, that pheasants and

INSTINCTIVE BEHAVIOUR OF YOUNG BIRDS. 89

partridges will scratch the ground, that a jay will go through
certain actions in the bath, that the preening of the down will
be carried out in particular ways—moor-hens, for example,
wringing out the water in a peculiar manner,—and that all these,
and many other modes of behaviour, will be presented in
relatively definite ways : all these are, to borrow the phrase of
Dr. Peckham’s, so characteristic of the several groups of birds,
that they would be an important part of any definition based
upon behaviour. And there can be no question that they are
instinctive. They may indeed seem trivial and commonplace,
scarcely worthy of special note; but they serve to show in
how many details organic heredity lays the foundation for
future behaviour, and affords groups of data for effective
consciousness to utilize.

To show the instinctive nature of such behaviour, the fol-
lowing examples will suffice. One of a batch of moor-hen
chicks showed once, and once only, when a week old, an in-
cipient tendency to bathe in the shallow tin of water which
was placed in their run, but soon desisted ; nor was the action
repeated, though he and the others enjoyed standing in the
water. Five weeks later one of the batch was taken to a beck.
He walked quietly through the comparatively still water near
the edge ; but when he reached the part of the stream where
it ran swiftly and broke over the pebbles, he stopped, ducked,
and took an elaborate bath, dipping his head well under,
flicking the water over his back, ruffling his feathers, and
behaving in a most characteristic manner. Hach day there-
after he did the same, with a vigour that increased up to the
third morning, and then remained constant. The same bird
some weeks later was swimming in a narrow part of the
stream, with steep banks on either side, when he was frightened
by a rough-haired pup. Down he dived, for the first time in
-his life; and after a few seconds his head was seen to appear,
just peeping above the water beneath the bank.

Ten days after receiving two nestling jays I placed in their
cage a shallow tin of water. They took no notice of it,
having never seen water before ; for they were fed chiefly on

go INSTINCTIVE BEHAVIOUR

sopped food. Presently one of them hopped into it, whether
attracted by the water or by accident it is difficult to say,
squatted in it bending his legs, and at once fluttered his feathers,
as such birds do when they bathe, though his breast scarcely
touched the water. The other seized the tin in his bill, and
then pecked at the water, thus wetting his beak. He, too,
fluttered his feathers in a similar fashion, though he was outside
the tin and not in the water at all. A little later the first
again entered the tin, and dipped his breast well into the water ;
this was followed by much fluttering and splashing. The
bird took a good bath, as did the other shortly afterwards,
and then spent half an hour in a thorough grooming, with
much fluttering of the wings, the crest feathers being con-
stantly raised and lowered, expressive of an emotional state.
Now, in these cases it would be impossible to say whether
the behaviour was carried out in the manner characteristic of
the species, prior to experience and independent of imitation,
on the basis of mere casual and chance observation. But in
these cases the whole life-history of the individuals concerned
was known; and it can be asserted with confidence that the
behaviour was hereditary, and not acquired by any gradual
process of learning. Moreover, in each case there seemed to
be such evidence as observation can afford, that internal
emotional factors co-operated with the direct external stimuli
in determining the nature of the behaviour. Whether such
actions so far contribute to the well-being of the individual as
to be of decisive advantage it is difficult to say. Some would
contend that bathing is practised by birds merely for the
pleasure it seemingly affords ; others would urge that it isa
means of getting rid of troublesome and presumably hurtful
parasites, to the attacks of which birds are peculiarly subject.
One of the most remarkable instincts of young birds is
that of the cuckoo, which ejects eggs and nestlings from the
home of its foster-parent. Mrs. Hugh Blackburn found a nest
which contained two meadow-pipits’ eggs, besides that of a
cuckoo. On a later visit “the pipits were found to be
hatched, but not the cuckoo. At the next visit, which was

INSTINCTIVE BEHAVIOUR OF YOUNG BIRDS 91

after an interval of forty-eight hours, we found the young
cuckoo alone in the nest, and both the young pipits lying
down the bank, about ten inches from the margin of the nest,
but quite lively after being warmed in the hand. They were
replaced in the nest beside the cuckoo, which struggled about

Fic. 17.—Young Cuckoo ejecting nestling Meadow Pipit. (From Mrs.
Hugh Blackburn's sketch in “ Birds from Moidart.”)

until it got its back under one of them, when it climbed back-
wards directly up the open side of the nest, and hitched the
pipit from its back on to the edge. It then stood quite upright
on its legs, which were straddled wide apart, with the claws
firmly fixed halfway down the inside of the nest, among the
interlacing fibres of which the nest was woven, and, stretching
its wings apart and backwards, it elbowed the pipit fairly over

92 INSTINCTIVE BEHAVIOUR

the margin, so far that its struggles took it down the bank
instead of back into the nest. As it was getting late, and the
cuckoo did not immediately set to work on the other nestling,
I replaced the ejected one and went home. On returning
next day, both nestlings were found dead and cold, out of the
nest.” * Here we have a definite account by an eye-witness,
who sketched the young cuckoo, which was naked, blind, and
could scarcely hold up its head. And her account, itself con-
firmatory of that given by Jenner in 1778, is confirmed by that
of Dr. John Hancock,t who witnessed the ejection of a fledg-
ling hedge-sparrow, which “was put over the edge of the nest
exactly as illustrated by Mrs. Blackburn.” The procedure is
unquestionably instinctive.

The sounds uttered by young birds are sufficiently definite
to be readily recognized and are susceptible of classification.
In domestic chicks at least six notes may be distinguished.
First the gentle “peeping ” note, expressive of contentment.
A further low note, a double sound, seems to indicate extremé
satisfaction and pleasure. Very characteristic and distinct is
the danger-note—a sound difficult to describe, but readily
recognized. If a humble-bee, a black-beetle, a big worm, a
lump of sugar-anything strange and largish—be thrown to
the chicks, this danger-note is at once heard ; and it serves
to place others on the alert, though this is perhaps the out-
come of experience. Then there is the cheeping sound,
expressive apparently of a state of mild dissatisfaction with the
present state of affairs. It generally ceases when one throws
some grain, or even stands near them. Extreme dissatisfaction
is marked by a sharper, shriller squeak, when one seizes them
against their inclination. Lastly, there is the shrill cry of
greater distress, when, for example, their swimming powers are
subjected to critical examination. With pheasants a gentle,
“peeping ” note of contentment, a shriller cry of distress, and
a danger-note, generically like, but specifically distinct from,

* «Birds from Moidart and Elsewhere,” p. 107. Edinburgh : Douglas.

+ Transactions of Northumberland and Durham Natural History Society,
vol. viii., p. 213,

INSTINCTIVE BEHAVIOUR OF YOUNG BIRDS 93

that of the chick, are early differentiated. The complaining
note of the partridge is uttered six or seven times in quick
succession, followed by a pause. The note of the plover is
high-pitched, and much like the familiar cry of the adult bird,
to which it owes its popular name of “ peewit.” So, too, the
guinea-fowl in down utters from the first notes quite charac-
teristic of its kind, while its danger-note is not unlike that of
the chick or pheasant. The piping of ducklings is compara-
tively nronotonous, and there does not seem to be a definite
danger-note. With moor-hen chicks, even on the first day,
two notes are well-marked—a call-note, lower in pitch than
that of the chick, and rather harsh and raucous, and a “tweet,
tweet” of pleasure, something like the contented note of a
canary. Later, five or six notes are differentiated, the most
characteristic of which is the harsh “crek, crek,” when the little
bird is from any cause excited. It is uttered in a crouching
attitude, with head thrown back and wings held outwards and
forwards, waving to and fro in a very characteristic manner.
That this has suggestive value for other moor-hen chicks is
shown by its distinctly infectious effect ; if one bird has cause
to utter the note and strike the attitude others follow suit.
While clearly instinctive in their mode of occurrence, while
they seem to show well the co-operation of an internal emo-
tional factor, their biological value seems to lie in their sug-
gestive effect on other members of the brood. They form an
elementary but sufficient social bond.

If these notes afford evidence of an incipient social factor,
the instinct of pecking is distinctively individualistic. Chicks
peck with considerable but not complete accuracy of aim at
practically anything of suitable size at suitable distance ; but
it is through experience that they learn what to select for food
and what to reject or leave untouched. Moving objects, how-
ever, are more readily pecked at than those which are still ;
and the instinctive response seems to be stimulated if one tap
on the ground near the object, or move it with a pencil, thus
simulating the action of the hen. And this is even more
marked with pheasants and partridges. Plovers seize small

94 INSTINCTIVE BEHAVIOUR

worms with an avidity which looks like an inherited recog-
nition of natural food. Pheasants and partridges also appear
to be specially affected by worms, and when one of them
seizes a worm for the first time, he shakes it and dashes it
against the ground. Chicks, a week or ten days old, also seize
a largish fly or bee with a dash, and maul it on the ground,
throwing it on one side before again approaching it. And
such birds seem to show an instinctive tendency to bolt with
such treasures as caterpillars or small worms. Moor-hens
cannot at first be induced to take food from the ground. It
has to be held above them, whereupon they crouch down, with
head and neck held back, opening their beaks more like the
callow young of nursling birds; but they also strike upwards
at the object—these modes of behaviour being, no doubt,
correlated with the manner in which the mother moor-hen
normally feeds her young from her beak during the early days
of life. Callow fledglings, such as young jays, simply open
their mouths, gaping widely to be fed. And many will
respond in this way to such a note as a low whistle, as may
readily be seen with swallows. But at a later age such birds
show instinctive modes of reaction of a more complex type.
A jay, for example, was offered a summer chafer or June bug,
seized it at once in his bill, and tried to place his foot on it.
Then he hopped down on to the floor of his cage, dropped the
beetle, seized it again as it crawled off, and after two or three
attempts swallowed it, tossing it back from the point of his
bill into the throat. This was the first time he took food
from the ground or swallowed it in this manner.

On the whole, there seems to be much inherited definiteness
of co-ordination, and some tendency to respond in a definite
manner to specific stimuli. That there should not be more
differentiation in this respect than observation discloses is
probably due to the fact that the parent birds afford, under
natural conditions, so much guidance in the selection of food.
Since the solitary wasp unerringly seizes its appropriate food,
since it responds instinctively to specific stimuli, there would
seem no reason why birds should not show similar instinctive

INSTINCTIVE BEHAVIOUR OF YOUNG BIRDS 95

differentiation. But one must remember that in the case of
the wasp there is no parental guidance; the insect is more
completely dependent on instinct than is the bird to whose
needs the hen assiduously ministers.

It is at first sight surprising that such birds as chicks and
pheasants do not peck instinctively at still water. When a
shallow vessel containing water was placed among some little
chicks, several of them ran repeatedly through the water, but
took no heed of it. Then, after about an hour, one of them
standing in the vessel pecked at his toes, and at once lifted
his head and drank freely with characteristic action. Another
subsequently pecked at a bubble near the edge, and then he
too drank. In fact, the best way of inducing them to drink
is to scatter some grains of food in the tin; they peck at the
grains, which catch their eye, and incidentally find the water,
and the touch of water in the bill at once leads to the cha-
racteristic response and congenitally definite behaviour. That
the sight of a still surface does not itself suffice to evoke this
behaviour is probably again due to the fact that under nature
the hen guides them and pecks at the water, when they follow
her bill.

One fact which must be constantly borne in mind is that
what is inherited is instinctive co-ordination, often related to
a definite stimulus, not instinctive knowledge. A chick pecks
at a grain when it is at a suitable distance, not because instinct
provides him with the knowledge that this is something to be
seized and tested, but because he cannot help doing so. He
is so organized that this stimulus produces that result through
an organic co-ordination that is independent of conscious
knowledge or experience. How definite is the inherited co-
ordination is shown by many observations. A young pheasant,
only a few hours old, was taken from the incubator drawer,
and held snugly while a piece of egg-yolk was moved before
his eyes with the aid of fine forceps. He did not peck at it,
but followed with movements of his head every motion of the
object in a narrow circle. Simple as this action seems, it
presents a striking example of co-ordinated movements

96 INSTINCTIVE BEHAVIOUR

accurately related to movements in the visual field, the whole
performed without any opportunity for learning or practice,
and less than half an hour after the bird was taken from the
drawer of the incubator and first saw daylight. Psychologists
sometimes puzzle their heads over the question how and by what
steps the field of vision and the field of movement are brought
into relation with each other; but in such a case as this, the
problem ceases to be primarily psychological. The relation is
purely organic ; the conscious data are grouped from the outset.
With young jays there was no such co-ordination at first ; and
when they began after a few days (about twelve or fourteen) to
follow an object with the head and eye, the movements were at
first jerky. But a week later, when I swept the food through a
circle a foot in diameter in front of their cage, it was followed
smoothly and evenly. Here a certain amount of learning and
practice, absent in the case of the pheasant, was required.
And it is difficult to say what proportion of the final result
was acquired, what proportion hereditary ; but probably the
behaviour is in the main instinctive, though somewhat
deferred.

One more example, perhaps even more trivial in the eyes
of some people, may be given. A duckling a few hours old
will scratch the side of his head. It is true he may topple
over in the process, through insufficient powers of balance, for
the simultaneous performance of poising on one leg and having
a good scratch with the other is no easy matter. But let not
either our familiarity with such behaviour, nor some observed
and laughable failure on the part of the duckling, blind us to
the fact that this is a congenital activity, and one of no little
complexity, indicating a definite organic nexus. A local irri-
tation sets agoing movements of the hind limb of that side
through which just that particular spot is scratched in the
absence of any previous practice, any learning to localize the
spot. There can be no question that such inherited co-ordi-
nations, whether perfect from the first, or with deferred per-
fection and some aid from acquisition, afford ready-made data
to consciousness, which are of the utmost service in the

INSTINCTIVE BEHAVIOUR OF YOUNG BIRDS 094

guidance of subsequent behaviour. The two-days-old chick,
with the aid of this instinctive co-ordination, performs well
a number of actions, which, had she to consciously learn them
all, would probably be still but half mastered when she was a
skinny old hen.

Our whole treatment of instinctive behaviour has been
based on the assumption, already to some extent justified, that
experience is not inherited. If it be hereditary, how comes it
that chicks show no recognition of still water, which must have
been familiar to the experience of generation after generation
of birds? How comes it that they do not even seem to
recognize their natural parent and protector, the hen? Two
chicks ten days old were taken to the yard whence were derived
the eggs from which they were hatched, and were placed about
two yards from a hen which was clucking to her brood. They
were not in a frightened condition, for they stood on my
hand and ate grain from it, scratching at the palm. But of
the clucking of the hen they took no notice whatever. The
same results were obtained with other chicks thirteen days old.
Was this due, as Spalding suggested, to loss of the instinctive
response which was perhaps present at an earlier age? Seem-
ingly not. For a chick was taken at the age of two and a half
days to its own mother, which had three chicks. These followed
her about, and ran at once to her when she clucked and pecked
on the ground. The little stranger, however, took no notice,
nor did he show any tendency either to go to the hen or to
follow the three chicks, having been purposely brought up
alone. When the hen took her little brood under her wing,
the stranger was placed close to her. She clucked, and seemed
anxious to entice and welcome the little fellow, seizing an oat-
husk and dropping it before him ; but he remained indifferent,
walking away and standing in the sunshine. After about forty
minutes he seemed more inclined to go with the other chicks,
but still ignored the existence of the hen. The natural
instinctive tendency seems to be from the first to nestle under
anything ; and there is the hen provided by nature for the
purpose. By experience the chicks grow accustomed to her fussy

H

98 INSTINCTIVE BEHAVIOUR

ways, as they grow accustomed to the ways of such a foster-
parent as the writer of these pages. Still, though there is,
apparently, no instinctive knowledge of the hen as their natural
protector, and though I have seen no observable response to
the clucking sound, this must not be taken as necessarily
implying that there is no instinctive response to any of her
modes of behaviour. There is such a response to her pecking
on the ground ; there is probably such a response to her danger-
note ; and there may be many other such instinctive modes of
behaviour related to her actions. How far they extend can
only be ascertained by patient observation ; and such responsive
behaviour need not imply any instinctive knowledge begotten.
of inherited experience.

We may now summarize some of the general conclusions
which may be drawn from observations of instinctive behaviour
in young birds.

1. That which is inherited is essentially a motor response or
train of such responses. Mr. Herbert Spencer’s description of
instinct as compound reflex action is thus justified.

2. These often show very accurate and nicely-adjusted
hereditary co-ordinations.

3. They are evoked by stimuli, the general type of which
is fairly definite, and may in some cases be in response to
particular objects.

+. They are also generally shown under conditions which
lead us to infer the “presence of an internal factor, emotional
or other.

5. There does not seem to be any evidence of inherited
knowledge or experience.

IV.—Tue Conscious Aspect or Instinctive BEHAVIOUR

In our definition of instinctive behaviour all positive refer-
ence to the presence of conscious states was omitted. By some
writers, however, the fact that it is accompanied by conscious-
ness is regarded asa distinguishing feature of instinct. Romanes

THE CONSCIOUS ASPECT 99

introduced his definition with the words : * “Instinct is reflex
action into which there is imported the element of conscious-
ness.” And he emphasized the conscious aspect when he said :
“The term comprises all those faculties of mind which are con-
cerned with conscious and adaptive action, antecedent to
individual experience.” Professor Wundt also lays some stress
on the conscious accompaniments of instinctive activities which,
he says,t “differ from the reflexes proper in this, that they are
accompanied by emotions in the mind, and that their perform-
ance is regulated by these emotions.” The definitions of other
writers express or imply the presence of consciousness in differ-
ing modes and degrees, culminating in the hypothesis of
inherited knowledge. Douglas Spalding, for example, said t
that “animals can forget the instinctive knowledge which they
never learned!”

Now, the exclusion from our definition of direct reference
to the conscious aspect must not be taken to imply that
instinctive behaviour is a mere matter of unconscious automa-
tism; nor even that it is unprofitable to discuss how much
consciousness there may be, of what sort, and how distributed.
All that it does imply is, that the amount, nature, and distribu-
tion of consciousness cannot well be introduced into a definition
the object of which is to help us to distinguish certain
observable types of behaviour from others. In a word, the
definition given is biological and objective, and is to be accepted
or rejected without prejudice to such psychological considera-
tions as those upon which we have now briefly to enter.

The first thing we have to decide is how much we are to
include, from the psychological standpoint, under instinct.
For we may take either a broader or a narrower view of the
matter ; and which of these we adopt will make much differ-
ence in our conclusions. Let us first deal with the narrower.
We have said above that what is hereditary in instinctive
behaviour is the co-ordination. Now, such co-ordination of

* “Mental Evolution in Animals,” p. 159.
+ “Lectures on Human and Animal Psychology,” p. 401.
+ “Instinct and Acquisition,” Nature, vol, xii., p. 507.

100 INSTINCTIVE BEHAVIOUR

movements into a finished and appropriate act is due to a
nicely graded distribution of efferent nerve-waves to the several
muscles concerned, so that these muscles may be caused to
contract in due order, and each to just the right extent. But
efferent nerve-waves as such, and their mode of distribution by
the nerve-centres, are in all probability unconscious, while the
contraction of the muscles is a purely organic matter. If,
therefore, we narrow our conception of instinct so as to include
only the co-ordinated act by itself, excluding all reference both
to the stimuli which are its antecedents, and to the effects in
consciousness which its performance may produce ; and if the
data for consciousness are in all cases supplied through
afferent channels ; then there seems to be no escape from the
conclusion that instinctive behaviour as such may be, and
probably is, altogether outside the individual consciousness.
It should be noted, however, that on this view it is the co-
ordination in itself that is, if not unconscious, at any rate
independent of the stream of experience.

Now, in the first place it is convenient so far to broaden our
conception as to include under the head of instinctive behaviour,
in its conscious aspect, not only the co-ordinated act but the
data which its performance affords to consciousness. It may
indeed seem that we are here trying to draw a distinction
where no real difference exists. The physiological distinction
is, however, not only clear and undeniable, but quite easily
understood. For the sake of illustration let us take the case
of an intentional action, such as glancing up from the words
we are reading to the clock. Efferent waves course along
several motor nerves to the six muscles by which each eye is
moved, and to the muscles of accommodation within the eye.
These muscles are called into duly co-ordinated activity, by
which our vision is focussed upon the clock-face. This is one
part of the physiological procedure—that by which the intended
result is attained. But there is a second part readily dis-
tinguishable from the former. As the eyes move, afferent
messages course inwards from the muscles or the eye-sockets and
their neighbourhood ; and it is these incoming waves which

THE CONSCIOUS ASPECT IOI

afford data to consciousness, telling us that the movements are
in progress or have been effected. The nerves involved in the
latter part are quite different from those concerned in the
former part, and they proceed to areas of the brain differently
situated from those whence the efferent waves issued. Thus
it is in all cases of movement; the efferent nerves call the
muscles into play; the afferent nerves bring information that
the movements are carried out. It is through the latter that
data are unquestionably afforded to consciousness.

But in the case of any complex action—and, as we have
seen, instinctive behaviour is often remarkably complex—the
information that the action has begun comes in before the
behaviour is completed. Practically we may say that any
given stage of performance and the consciousness it evokes are
simultaneous ; for though in strictness the one lags just a
little behind the other, yet they are so nearly coincident in
time that we may disregard the interval between them. Such
being the case, therefore, we may fairly regard instinctive
behaviour as capable of introducing important elements into
the conscious situation.

But not only does instinctive behaviour thus introduce
important elements into the conscious situation, it is also
called forth by stimuli which themselves afford not less
important elements. To exclude these from any consideration
of instinct, in its conscious aspect, would render the treatment
of the problem so incomplete as to be wholly unsatisfactory
from a psychological point of view. Can we believe that
when the moor-hen dived, as it never had dived before, at the
sight of the rough-haired pup, the vivid experience of that
strange and disquieting intruder did not enter into, and form
a prominent feature in, the conscious situation ? If we are to
consider the conscious aspect at all, we must try and grasp the
situation as a whole. And on these grounds we may yet
further broaden our conception so as to include, from the
psychological point of view, not only the behaviour itself, and
its effects in consciousness, but also the conditions under which
the complex actions are called into play. If, then, we accept

102 INSTINCTIVE BEHAVIOUR

this position, and agree to use the term “instinct” for our
present purpose in a comprehensive sense, we may now proceed
to consider very briefly the nature of the elements which enter
into the instinctive situation.

First, there are the external stimuli affecting one or more
of the sense organs, and thus evoking consciousness ; and
secondly, there are internal factors, having their source in the
condition of the body, or its parts and organs. It is con-
venient to take these two together, so that we may see what
relationship they bear to each other. Both seem to be present,
and to co-operate in a great number of instinctive acts. In
the behaviour connected with feeding, for example, an internal
element of hunger co-operates with the external presentation
of the appropriate food or prey. So, too, with the instincts
concerned in the propagation of the race. Looking at the
matter generally, we may regard the internal factors of the
kind with which we are now dealing, as giving rise to a want
or need, passing in some cases into a state of craving. In
themselves such conscious states are in their inception ex-
ceedingly indefinite ; for a want can only be rendered definite
in experience by its appropriate satisfaction. In many cases
of instinctive behaviour the indefinite want and the particular
and duly related stimulus seem to lead, without prevision and
by a blind impulse, to the performance of those acts which
will afford the unforeseen satisfaction. And when once this
satisfaction has been attained, subsequent wants or needs of
like character will no longer be indefinite ; nor will future
behaviour of the same kind be thereafter wholly instinctive,
for it can never again be prior to, or independent of, experience.

Granted, however, that a felt need of some kind, indefinite
at first but none the less real, is present in many cases as a
spur to instinctive behaviour ; is it in all cases a necessary
factor ? May we say that this distinguishes instinctive from
merely reflex action ? The question is, from the nature of the
case, exceedingly difficult to answer. But without going so
far as to say that reflex action may be unerringly distinguished
from instinctive behaviour by the absence of any such internal

THE CONSCIOUS ASPECT 103

factors, we may perhaps, at any rate, go so far as to give pro-
visional acceptance to the view that in instinct these wants
and felt needs enter into the conscious situation in a manner
and to a degree that are so far distinctive—which seems to be
the position adopted by Professor Wundt.

There is, however, a further relation between the external
stimulus and these internal factors which is presumably of no
little importance. The stimulus intensifies the want, or may
in some cases call it into existence. Just as a whiff from the
kitchen may lead us to realize that we need the satisfaction
that will erelong be presented at table, so may the sight of
his mate in the spring evoke in the breast of the yearling
sparrow a need, having its source in morphological and
physiological changes, that spurs him on to the courtship that
shall lead to its due satisfaction. Popular attention has,
indeed, been so naturally drawn to the internal needs or wants
with which we are now dealing, as to give them an almost
exclusive monopoly of the term “ instinct,” which thus often
comes to be regarded asa connecting link between the stimulus
and the act. The sight of a mouse, for example, is said to call
forth the instinct of the cat, which is satisfied by her pouncing
upon it. And so it comes about that, while the biologist
fixes upon the instinctive act as the essential feature, the
psychologist is apt to regard the impulse * which prompts to
action as the more central and characteristic element. We are
here endeavouring to combine both these points of view.

To come to closer quarters with the relationship which
holds good between the external and internal elements, it
appears that, when the stimulus evokes or intensifies the want
or need, this is probably effected by efferent waves which call
the organs or parts into tonic action, of which the animal
becomes conscious through the afferent messages which come
in from them to the sensory centres ; in much the same way
as the whiff from the kitchen takes effect on the salivary and
other glands, and throws the organs of digestion into a felt
preparedness for the fulfilment of their functions. But it

* On the nature of impulse, see infra, p. 235.

104 INSTINCTIVE BEHAVIOUR

may have other and more indirect consequences. When the
moor-hen dived to escape from the obtrusive puppy, his heart-
beat was probably affected ; he had, perhaps, an uncomfortable
sinking in his gizzard ; his breathing was short and laboured ;

and he experienced creepy sensations in the skin and around... .

the feather-roots. Such we may suppose were the accompani-
ments or sources of the emotional state of fear or alarm. And
they presumably entered with no little vividness into the con-
scious situation at the moment of instinctive action. In all
those cases in which the behaviour is associated with such an
emotional state as anger or fear, the external stimulus seems to
produce widely-spread effects on the glands, respiratory organs,
heart and blood-vessels, skin and other parts, as well as the
more direct response in productive action. And all this must
enter into the conscious situation, contributing largely, as we
shall hereafter see, to the emotions in their instinctive origin.

Enough has now been said to indicate with sufficient clear-
ness the kind of co-operation and mutual relationship which
subsists between the external and the internal factors in the
conscious situation which leads to instinctive behaviour. We
have seen that, not improbably, some organic prompting is
always present in greater or less degree. But the question
still remains whether anything like a definite and particular
external stimulus is in all cases a necessary factor.

When the predaceous larva of the water-beetle, Dytiscus,
ceases to feed, and, creeping into the moist earth near the
pond’s edge, makes a hollow cell in which to enter upon its
pupal sleep, there does not seem to be any well-defined stimulus
from the outer world which can be said to initiate the be-
haviour of whose purport the larva can have no idea. Some
inner need seems to impel the creature to this necessary but as
yet unknown course of action ; and this appears to constitute,
if not the sole, at least the preponderant element in the
conscious situation. In healthy young birds and other animals
there is after the rest of sleep a certain exhilaration and
exuberance of spirits which seemingly leads to characteristic
action ; dancing, flapping of the wings, running hither and

THE CONSCIOUS ASPECT 105

thither in short quick spurts, and so forth. No doubt in such
cases external stimuli are present, and contribute in some
degree to the effects produced; but they do not seem to be
particularized so that one can say that just this or that well-
defined stimulus is necessary to give rise to the observed
behaviour. In the case of migration, too, an internal factor—
the nature of which we do not know—is probably as strong
as if not stronger than any influence from without. While,
therefore, we may say that some external factors are frequently,
not improbably always, contributory, we must add that observa-
tion does not enable us in all cases to define them with any
approach to accuracy; and, further, that promptings from
within seem in some instinctive acts to be the most important
elements in the conscious situation.

It now only remains to draw attention to the fact that the
effects of the behaviour, as the animal becomes conscious of
the performance of the acts concerned, serve to complete and
render definite the conscious situation. Consciousness, how-
ever, probably receives information of the net results of the
progress of behaviour, and not of the minute and separate
details of muscular contraction. These net results, having
thus entered presentatively into the situation, are subsequently
susceptible of re-presentative recall, when the recurrence of
certain salient elements serve to reproduce the essential features
of the situation of which experience has been gained on a
former occasion. Hence, as has already been noted, it is only
the first performance of an instinctive action which can be
described as prior to experience. The second time the deed is
done it is done by an animal which has had opportunity of
gaining experience on the foregoing occasion. And then it
may be done with a difference, with some acquired modification
of performance. By the repetition of the slightly modified be-
haviour the effects of habit are introduced, and thus acquired
peculiarities of action are established as individual traits. We
must not forget that, in a large number of cases, so-called
instinctive behaviour, as presented to observation, has lost
through modified repetition its original purity of type. The

106 INSTINCTIVE BEHAVIOUR

acts we see are often the joint products of heredity and in-
dividual acquisition, the inherited co-ordination having been
supplemented or otherwise altered through experience.

Even in the case of the very first exhibition of such a deferred
instinct as the moor-hen’s dive, although that organized
sequence of acts which constituted the behaviour as a whole
had never before occurred, although there was no gradual
learning how to dip beneath the surface, and to swim under
water, still many of the constituent acts had been. often
repeated ; experience had already been gained of much of the
detail then for the first time combined in an instinctive
sequence. So that if we distinguish between instinct as con-
genital and habit as acquired, we must not lose sight of the
fact that there is continual interaction, in a great number of
cases, between instinct and habit, and that the first performance
of a deferred instinct may be carried out in close and in-
extricable association with the habits which, at the period of
life in question, have already been acquired. Instinct supplies
an outline sketch of behaviour, to which experience adds colour
and shading. Which predominates in the finished picture
depends on the status of the animal. In the lower and less
intelligent types the outline stands out clearly, there being but
little shading to divert our attention from the clear firm lines
inscribed by heredity ; but in the higher and more intelligent
animals, the deft pencil of experience has added so much and
has interwoven with the fainter outline so many new and
skilfully introduced touches, that the original sketch is scarcely
distinguishable unless we have carefully watched from the
beginning the gradual development of the picture.

V.—Tue Evouvution or INstincTIvE BEHAVIOUR

We have seen that Professor Wundt distinguishes two
classes of instinctive acts : first, those which are acquired or
have become wholly or partly mechanized in the course of
individual life ; secondly, those which are connate or have been
mechanized in the course of generic evolution. ‘The laws of

EVOLUTION OF INSTINCTIVE BEHAVIOUR 107

practice,” he says,* “suffice for the explanation of the acquired
instincts. The occurrence of connate instincts renders a sub-
sidiary hypothesis necessary. We must suppose that the
physical changes which the nervous elements undergo can be
transmitted from father to son. . . . The assumption of the
inheritance of acquired dispositions or tendencies is inevitable
if there is to be any continuity of evolution at all. We may
be in doubt as to the extent of this inheritance; we cannot
* question the fact itself.”

Now, the application of the term “instinct,” both to
acquired and to connate behaviour, seems to prejudge the
question of their genetic connection. And since we have the
well-recognized term habits for actions the performance of
which becomes automatic through frequency of repetition, we
may substitute this term, or the phrase habitual acts, for the
“acquired instincts” of Professor Wundt. Modifying, there-
fore, his statement in accordance with this usage, the fact
which, he says, we cannot question is that acquired habits are
inherited as congenital instincts. This opinion has long been
held: G. H. Lewes regarded instinctive actions as transmitted
habits from which the intelligence, through which they were
originally acquired, had lapsed. Darwin believed that such
inheritance was a factor in the evolution of instinctive be-
haviour. Romanes distinguished instincts due to this mode of
origin as “secondary ;” reserving the term “primary” for
those attributable to natural selection, and describing those in
which both factors co-operate as “ instincts of blended origin.”
The late Professor Himer, of Tiibingen, going further than
either Darwin or Romanes, reverted almost entirely to what we
may term the Lamarckian interpretation. “I describe as
automatic actions,” he says,t “those which, originally per-
formed consciously and voluntarily, in consequence of frequent
practice come to be performed unconsciously and involun-
tarily. . . . Such acquired automatic actions can be inherited.
Instinct is inherited faculty, especially is inherited habit.” In

* “Lectures on Human and Animal Psychology,” p. 405.
+ “ Organic Evolution,” pp. 223, 263, 258, 279, 276, 298.

108 INSTINCTIVE BEHAVIOUR

his discussion of the subject Eimer makes no express allusion
to primary instincts ; but he attributes to lapsed intelligence
some of those which were classed by Romanes as primary, and
his tendency is to refer all instincts to the same source.
“ Every bird,” he says, ‘“‘ must, from the first time it hatches its
eggs, draw the conclusion that young will also be produced
from the eggs which it lays afterwards, and this experience
must have been inherited as instinct.” Why, in the first
instance, it must draw the conclusion from observation if it
inherit instinctive knowledge, is not made clear. But our
present purpose is to indicate, not to criticize, Himer’s position.
He claims that “‘the original progenitors of the cuckoo, when
they began to lay their eggs in other nests, acted by reflection
and design.” Of the behaviour of mason wasps and their allies,
which provide their young with paralyzed but living prey, he
exclaims, “ What a wonderful contrivance! What calculation
on the part of the animal must have been necessary to discover
it!’ Of the instincts of neuter bees he remarks, “ Selection
cannot here have had much influence, since the workers do not
reproduce. In order to make these favourable conditions
constant, insight and reflection on the part of the animals, and
the inheritance of these faculties were necessary.” And he
concludes, ‘‘ Thus, according to the preceding considerations,
automatic action may be described as habitual voluntary
action ; instinct, as inherited habitual voluntary action, or
the capacity for such action.”

Turning now to the opposite end of the scale of opinion,
we find that Professor Weismann, commenting on the supposed
inheritance of acquired habit, says,* “I believe that this is an
entirely erroneous view, and I hold that all instinct is entirely
due to the operation of natural selection, and has its founda-
tion, not upon inherited experiences, but upon variation of
the germ.” Ziegler and Groos in Germany, Whitman and
Baldwin in America, Poulton and Wallace in England, either
deny the existence of secondary instincts, due to'the inheritance
of acquired habits, or question the sufficiency of the evidence

* «Essays on Heredity ” (1889), p. 91.

EVOLUTION OF INSTINCTIVE BEHAVIOUR 109

adduced in support of such transmission. In their explanation
of the manner in which that inherited co-ordination, which is
biologically the central fact in instinctive behaviour, has been
evolved they rely mainly or entirely on the principle of natural
selection.

What, then, were the facts which appeared to Romanes
sufficient to justify a belief in the existence of a class of
instincts dependent on inherited habit for their origin? He
tells us that he only gives a few examples ‘“‘ amongst’ almost
any number” that he could quote. It is certainly unfortunate
that, out of more than one hundred and fifty pages devoted to
instinct in his work on ‘“‘ Mental Evolution in Animals,” only
three * are assigned to secondary instincts ; or six, if we include
one dealing with inherited peculiarities of hand-writing in
man, and two showing the force of heredity in the domain
of instinct, “ whether of the primary or secondary class.” It
is true that many pages are devoted to instincts of blended
origin, but the co-operation of the Lamarckian factor is here
rather assumed than proved. We must, however, be content
to take the few examples that are actually given. They are
four in number. First, that ponies in Norway are used with-
out bridles, and are trained to obey the voice; and that, as
a consequence, a race-peculiarity has been established, for
Andrew Knight says that it is impossible to give them what
is called “a mouth.” No details being given, this strikes one
as rather thin as a matter of evidence. Secondly, Mr. Lawson
Tait had a cat which was taught to beg for food like a terrier.
All her kittens adopted the same habit under circumstances
which precluded the possibility of imitation. Supposing the
facts to be correctly reported, and granting that the owners
of the kittens, presumably aware of the maternal propensity,
did not take some pains to teach the offspring of such a parent
to beg (and this does not present much difficulty), one can
hardly found a scientific conclusion on so slight an anecdotal
basis. ‘Thirdly, instinctive fear is said to be an inherited
acquisition ; which, fourthly, is lost by disuse. But, as we

* Op. cit., pp. 196-198.

IIo , INSTINCTIVE BEHAVIOUR

have already seen, modern investigation has placed this matter
of so-called hereditary fear of natural enemies on a different
footing. Pheasants, partridges, moor-hens, and wild duck show
no fear of a quiet dog. If approached gently, in the absence
of their parents, callow wild birds in their nest exhibit little
alarm at the slow and gentle approach of man. Mr. Hudson’s
opinion has already been quoted, but will bear repetition ; it
is, “that fear of particular enemies is in nearly all cases the
result of experience.” And there is no evidence to show that,
in those cases in which it is truly instinctive and not the result
of experience, the instinctive behaviour is necessarily due to
inherited habit and not to natural selection.

It cannot be said that the evidence for the supposed mode
of origin of secondary instincts is sufficiently varied and cogent
to carry conviction. On the other hand, there does seem
some evidence which points to a different conclusion. When
instinctive behaviour follows on a sensory impression, not only
is the co-ordination hereditary, but there is an inherited link-
age of stimulus and response. Thus in the solitary wasps the
sight of the natural prey is followed by the appropriate modes
of attack. The J/e/oé larva springs upon anything hairy. In
chicks the sight of a small object at a certain distance initiates
the act of pecking. In moor-hens and ducklings the stimulus
of water produces the movements concerned in swimming.
And so, too, in many other examples of instinctive behaviour,
we infer from the observed facts that stimulus and response
have an organic connection founded on hereditary links in the
nervoussystem. Now, if such connection were due to inherited
habit, we should expect them to be established wherever the
experience to which they are related has been constant through
wavy generations. How comes it, then, that the chick does
not instinctively respond by appropriate behaviour to the
sight of water? How comes it that young birds do not
instinctively avoid bees, and wasps, and nauseous caterpillars ?
If the effects of ancestral experience be hereditary, one would
surely expect that in these cases the connection between
stimulus and response—a connection which passes into acquired

EVOLUTION OF INSTINCTIVE BEHAVIOUR 111

habit—would have become congenital; that the habitual
behaviour would have long ago become instinctive. But this
does not appear to be the case. And with regard to disuse
causing the loss of instinct, how comes it that young chicks
swim with well-ordered leg-movements, though swimming is
not an act that is habitually performed by the members of
their race ?

What, then, has the alternative hypothesis of natural
selection to advance in explanation of these facts? On this
hypothesis instinctive acts have biological value in such degree
that they have become congenital through the preservation of
adaptive variations. But if this be so, why does not the chick
respond instinctively to the sight of that which is so essential
to its existence as water to drink? In reply to this question
it may be suggested that, under natural conditions, the hen
teaches all her chickens to peck at the water, and thus shields
them from the eliminating influence which gives rise to natural
selection, in the absence of which the habit of drinking in
response to the sight of water, though acquired by each
succeeding generation of birds, has not become instinctive and
congenital. Or, to put the matter from a slightly different
point of view, the maternal instincts of the hen protect her
chicks from any elimination in this respect ; and in the absence
of such elimination the habit has not been inherited as instinct.
But though the hen can lead her young to peck at the water,
she cannot teach them how to perform the complex move-
ments of the mouth, throat, and head in actual drinking. In
this matter, therefore, her own instinctive procedure does not
shield them from the incidence of that elimination which leads
to survival under natural selection. Those chicks which, on
pecking the water, failed to respond to the stimulus by the
complex behaviour involved in drinking would be eliminated,
leaving those to survive in which the response had been
congenitally established. Thus it would seem that, when
natural selection is excluded, the habit has not become con-
genitally linked with a visual stimulus; but when natural
selection is in operation, the response has been thus linked

112 INSTINCTIVE BEHAVIOUR

with the stimulus of water in the bill. Whence we may infer
that the co-operation of natural selection is an essential factor _
in the evolution of instinctive behaviour.

There are, however, cases of instinctive behaviour which
may seem too trivial and unimportant to be subject to the
sway of natural selection. There are numberless little idio-
syncracies of behaviour which seem to be truly instinctive,
which are readily recognizable as distinctive traits, but which
can hardly be regarded as of sufficient biological value to
determine whether the creatures in which they are developed
should survive or be eliminated in the struggle for existence.
In many cases, however, these serve rather to distinguish the
detailed manner of behaviour than its biological end or
purpose. In different species natural selection may determine
the survival of those whose instinctive behaviour meets a
biological need. The relatively unimportant details, differing
slightly in each species, are mere adjuncts ; and since natural
selection deals with each species or inter-generating group
separately, the essential behaviour may in each case carry with
it the associated differences of manner. We must remember,
too, that, as in the matter of structure so in that of behaviour,
it is the animal as a whole that is selected for survival ; and
so long as the whole is adapted to the circumstances of life,
the associated differences of form or manner may share in,
without doing much to determine, survival. In any case these
little instinctive traits, if they are so trivial as to seem of small
value from the biological point of view, appear to be too
unimportant to have been intelligently acquired as habits.

Let us now consider one or two cases of instinctive behaviour
which would fall under Romanes’s category of instincts of
blended origin partly due to natural selection, partly to the
inheritance of acquired habit. It is the custom of the house
martin to build beneath the eaves. Forsaking the ancestral
rocky haunts, it has been led to utilize the houses that
man has built. This has all the appearance of being due
to an intelligent modification of the ancestral instinct ; but
how far the modification has become through heredity a

EVOLUTION OF INSTINCTIVE BEHAVIOUR 113

congenital variation, we do not know. The intelligence
which is said to have enabled the martin of the past to adopt
this method of nidification is still operative. The nestlings
brought up under the eaves would have opportunities for
acquiring experience which might lead them to build under
similar circumstances. Nest and eaves would be associated in
the conscious situation. Nor would the effects of natural
selection be necessarily excluded. One may suppose that in
the open country, far from rock-shelters, those martins in
which there was a congenital tendency to build in house-
shelters would bring up their broods and transmit this
tendency ; while those in which it was absent would either go
elsewhere or fail to bring up broods at all. In the absence
of fuller knowledge as to the truly instinctive nature of the
behaviour, and as to its mode of genesis, we are in large
degree at the mercy of conjecture. But in any case the in-
cidence of elimination is not necessarily excluded, and there
are, therefore, no grounds for denying that natural selection
has been a co-operating factor in the evolution of the instinc-
tive behaviour, if such it be.

It is well known that the lapwing will apparently simulate
the actions of a wounded bird, with the object, as it seems, of
drawing intruders away from her nest. And such tactics are
not restricted to this bird, nor even to one or two species.
They are common, no doubt with diversities of detail, to such
different birds as grouse, pigeons, plovers, rails, avocets, ducks,
pipits, buntings, and warblers. Granting that the behaviour
is truly instinctive, it forms a very pretty subject for trans-
missionists and their critics to quarrel over. “If we seek, as
an-example,” the transmissionist may exclaim, “an instinct
which bears the marks of its intelligent, and therefore acquired
origin, this of feigning wounded provides all that we can
possibly demand.” “What mode of instinctive behaviour,” the
selectionist may ask, ‘can be adduced which is more obviously
useful to the species? Is not this just the kind of procedure
which natural selection, if it be a factor at all, must fix upon
and perpetuate through the elimination of failures? Those

1

114 INSTINCTIVE BEHAVIOUR

birds which, through congenital variation of behaviour, acted
in this way would certainly enable their offspring to escape
destruction by enemies, and these would survive to perpetuate
the instinct.”

Let us expand the transmissionist position a little further.
An extremist, of the type presented by Eimer, would perhaps
urge that the lapwing reasons thus: “If I pretend to be
wounded, trail my wing, and flutter along the ground, instead
of flying off, I shall draw upon myself the intruder’s attention,
and lead him to suppose that I shall be easily caught ; and if
I thus entice him away, my little ones will be saved, and my
end gained.” Thus, it may be said, might the bird argue,
and then give practical effect to its reasoning. But are we
not here attributing to the lapwing powers of ratiocination
beyond the capacity of the most intelligent of birds? Are
we not assuming a histrionic power, and a realization of the
effects on others of its display, which many a human actor
might well covet ?

“But may not the bird,” it may be urged in reply, “have
found by experience, without any elaborate process of abstract
reasoning, that the trick is effectual?” In any case it would
be experience perilously acquired. Granting that the bird has
the wit to try the trick, a little over-acting, a little too much
lameness of wing, and she is herself seized and killed ; a little
under-acting, and the trick fails—her brood is found and
destroyed. Does it not seem probable that such experience
would be dearly bought, that failure would mean either death
to the parent or death to the offspring? And is it not clear
that natural selection is thus introduced in any case? And
may not the selectionist pertinently ask: “ Why, if natural
selection is thus introduced as a factor, halt midway between
two hypotheses? Why not take the further step—one by
which all the difficulties attending the intelligent acquisition
and the biological transmission are alike avoided—of allowing
that natural selection exercises, throughout, its influence on
congenital variations, and not on acquired modifications of
behaviour ?”

EVOLUTION OF INSTINCTIVE. BEHAVIOUR 115

There is, however, a way in which, when natural selection
is operative, intelligence may serve to foster congenital varia-
tions of the required nature and direction. We must re-
member that acquired habits on the one hand, and congenital
variations of instinctive behaviour on the other hand, are both
working, in their different spheres, towards the same end, that
of adjustment to the conditions of life. If, then, acquired
accommodation and congenital adaptation reach this end by
different methods, survival may be best secured by their co-
operation. And the more thorough-going the co-operation
the better the chance of survival. There would be a dis-
tinct advantage in the struggle for existence when inherited
tendencies of independent origin coincided in direction with
acquired modifications of behaviour ; a distinct disadvantage
when such inherited tendencies were of such a character as to
thwart or divert the action of intelligence. Thus any here-
ditary variations which coincide in direction with modifications
of behaviour due to acquired habit would be favoured and
fostered ; while such variations as occurred on other and
divergent lines would tend to be weeded out. Professor Mark
Baldwin,* who has independently suggested such relation
between modification and variation, has applied to the process
the term “Organic Selection ;” but it may also be described
as the natural selection of coincident variations.

It may be urged, therefore, that if natural selection be
accepted as a potent factor in organic evolution, and unless good
cases can be adduced in which natural selection can play no
part and yet habit has become instinctive, we may adopt some
such view as the foregoing. While still believing that there is
some connection between habit and instinct, we may regard the
connection as indirect and permissive rather than direct and
transmissive. We may look upon some habits as the acquired
modifications which foster those variations which are coincident
in direction, and which go to the making of instinct.

The net result of a study of instinctive behaviour is to lead

* Professor Henry Osborn has also indicated the relationship re-
ferred to. ca

116 INSTINCTIVE BEHAVIOUR

us to the conclusion that its evolution runs parallel with the
evolution of animal structure. This is perhaps best seen in
the case of those insects in which typical instinctive acts are
performed by larvee of wholly different form and structure,
though they are stages in the development of the same species.
This is exemplified in the cases of Sitaris, Argyromeba, and
Lewopsis which have, been briefly described. It is probable
that in all cases of instinctive action natural selection has been
a co-operating factor. Without going so far as to assert with
Professor Weismann the “all-sufficiency of natural selection,”
we may echo the words of Professor Groos,* and say : “ Never-
theless, we know no principle except that of selection, and we
must go as far as that will take us. Absolute knowledge of
‘such phenomena is unattainable.” And in this conclusion we
have the support of Dr. Peckham, who says,f “ We have found
them [the instinctive acts of solitary wasps] in all stages of
their development, and are convinced that they have passed
through many degrees from the simple to the complex, by the
action of natural selection. Indeed, we find in them beautiful
examples of the survival of the fittest.”

* «The Play of Animals,”’ p. 64.
t “Solitary Wasps,” p. 236.

CHAPTER IV

INTELLIGENT BEHAVIOUR

I.—Tuer Nature or Inretiicent BEHAVIOUR

SucH an animal as a newly hatched bird or an insect just set
free from the chrysalis is a going concern, a living creature.
It is the bearer of wonderfully complex automatic machinery,
capable, under the initiating influence of stimuli, of performing
instinctive acts. But if this were all we should have no more
than a cunningly wrought and self-developing automatic
machine. What the creature does instinctively at first it
would do always, perhaps a little more smoothly as the organic
mechanism settled down to its work—just as a steam-engine
goes more smoothly when it has been running for a while ;
but otherwise the action would continue unchanged. Instinc-
tive behaviour would remain unmodified throughout life. The
chick, however, or the imago insect is something more than
this. It affords evidence of the accommodation of behaviour
to varying circumstances. It so acts as to lead us to infer
that there are centres of intelligent control through the
action of which the automatic behaviour can be modified in
accordance with the results of experience. When, for example,
a young chick walks towards and pecks at a ladybird, the like
of which he has never before seen, the behaviour may be purely
instinctive ; and so, too, when he similarly seizes a wasp-larva.
Even when he rejects the ladybird or swallows the larva, this
may be directly due to unpleasant stimulation in the one case,
and pleasant in the other. But when, after a few trials, the
chick leaves ladybirds unmolested while he seizes wasp-larve

117

118 INTELLIGENT BEHAVIOUR

with increased energy, he affords evidence of selection based
on individual experience. And such selection implies intelli-
gence in almost its simplest expression. We may say, there-
fore, that, whereas instinctive behaviour is prior to individual
experience, intelligent behaviour is the outcome and product
of such experience. This distinction is presumably clear
enough ; and it is one that is based on the facts of observa-
tion. But we must not fail to notice that, though the logical
distinction is quite clear, the acquired modifications of be-
haviour, which we speak of as intelligent, presuppose congenital
modes of response which are guided to finer issues. We may
say, then, that where these congenital modes of response take
the form of instinctive behaviour, there is supplied a general
plan of action which intelligence particularizes in such a
manner as to produce accommodation to the conditions of
existence.

We have already frankly admitted that, in the present state
of our knowledge, we do not know with any definiteness how
intelligent modification of behaviour is effected. But it seems
probable that from all parts of the automatically working
organic machine messages come in to the centres of conscious
control, and that in accordance with the net result of all these
messages, past and present, tinged with pleasure or pain, other
messages go out to the automatic centres, and, by checking
their action here and enforcing it there, give new direction to
the resulting behaviour. If this be so, the consciousness
associated with the control-centres is like the person who sits
in a central office and guides the working of some organized
system in accordance with the information he is constantly
receiving ; who sends messages to check activity in certain
directions and to render it more efficient and vigorous in
others.

It may be said, however, that intelligent guidance is, at any
rate in such simple cases as the selection of a palatable grub
and the rejection of a nauseous ladybird, itself determined by
instinctive likes and dislikes. All young chicks apparently
find wasp-larve palatable and ladybirds the reverse; and

THE NATURE OF INTELLIGENT BEHAVIOUR 119

this is just as much the outcome of heredity as the instinc-
tive act of pecking. Since, therefore, heredity determines
what shall be selected and what rejected—since the likes and
dislikes are themselves instinctive—any essential difference
between congenital and acquired behaviour seems to be
evanescent.

Now, if we apply to the affective qualities of mental states
—the pleasurable tone or its opposite which characterize such
states—the term “instinctive,” we do so in reference to the
broader psychological conception of instinct, rather than in
accordance with the narrower biological acceptation of the
term. For the likes and dislikes constitute part of the con-
ditions under which the behaviour occurs, and not elements in
the co-ordinated response as such. Hence it is preferable to
apply to these hereditary qualities the term inate, rather than
the term instinctive. But, waiving this distinction, it is true
that such pleasant or unpleasant qualities of the sensory results
of stimuli are part of the animal’s hereditary outfit, and are
not acquired in the course of individual experience. What,
then, is acquired? What part does experience play in the
development of intelligent behaviour? Let us consider the
case of the chick and the ladybird, and see whether it helps
us to answer these questions. The chick is stimulated to the
instinctive pecking response by a small moving object. That
is the first scene of the little drama. In the second scene the
ladybird is seized, sensory centres are unpleasantly stimulated,
and the insect is dropped or thrown on one side with signs of
disgust. Let us grant that this aversion with its characteristic
response is also instinctive. There is no hereditary connection
between scene 1 and scene 2. After an interval the curtain
rises on act ii. The characters are the same as in the first
scene of the previous act. But the action of the drama is
different. The chick does not seize the ladybird. Why?
Because there is an acquired connection between scenes 1 and 2
of the previous act. The chick has gained experience of the
nauseous character of the insect, and this experience influences
and modifies his behaviour. The essentially new feature,

120 INTELLIGENT BEHAVIOUR

therefore, is the establishment of a connection which is not
provided through inheritance. To put the distinction in a
brief form, we may say that instinct depends on how the
nervous system is built through heredity ; while intelligence
depends upon how the nervous system is developed through
use.

Assuming that an animal is capable of gaining experience
and of acquiring new nervous connections in the course of
individnal experience, it follows that, as has already been
indicated, instinctive behaviour in its logical purity is only
presented in the first performance of any given co-ordinated
act. For after this the animal has gained experience of its
performance ; and this can no longer conform to a definition
of instinct, according to which it is characterized as “ prior to
experience.” On the other hand, intelligent behaviour cannot
be presented on the first occurrence of any action, since there
is no prior experience thereof in the light of which it may be
guided. This logical distinction may be expressed by saying
that instinctive behaviour is always prior to experience, while
intelligent behaviour is always subsequent to experience.
When, however, instinctive procedure continues throughout
life practically unmodified or but little modified, we may still
class it under instinct, since the hereditary connections are
still the predominant factors. And where the latter part of
an instinctive sequence is modified by the experience gained in
the former part, we may still term the modification intelligent,
however small may be the time-interval implied in the word
“subsequent.” Sharp as the logical distinction is, the be-
haviour of animals is in the main a joint-product, and whether
we term it instinctive or intelligent depends upon whether the
hereditary or the acquired factor predominates.

Passing on now to consider some further characteristics of
intelligent behaviour, we may first notice what Dr. Charles
Mercier, in his work on ‘‘ The Nervous System and the Mind,”
terms the four criteria of intelligence. Intelligence is mani-
fested, he says, first in the novelty of the adjustments to
external circumstances ; secondly, in the complexity ; thirdly,

THE NATURE OF INTELLIGENT BEHAVIOUR 121

in the precision ; and fourthly, in dealing with the circum-
stances in such a way as to extract from them the maximum
of benefit.

Tf, however, we are to regard these severally as criteria of
intelligence, each should serve to differentiate intelligent from
instinctive behaviour. But this is not the case. The precision
of the adjustment can-
aot be regarded as a cri-
terion of intelligence, for
many instinctive acts are
remarkably precise. No
grocer’s assistant rolls a
paper funnel with more
precision than is dis-
played by the birch-wee-
vil (Rhynchites betule) in
constructing the leaf-
case in which her eggs
are laid. Curved inci-
sions of constant form
are made on either side
of the midrib, and are
“of just the right shape
to make the overlaps in
the rolling, and to re-
tain them rolled up
with the least tendency
to spring back,” * while the tip of the leaf is rolled into
a second smaller funnel, which is tucked in to close the
opening of the first, after the eggs have been deposited. “The
eggs hatch in their dark place, each giving rise to an eyeless
maggot, which ultimately leaves the funnel for the earth. . . .
Hence the beetle cannot be considered to have ever seen a
funnel, and certainly has never witnessed the construction of

1

Fig. 18.—Leaf-case of Birch-weevil.

* See D. Sharp, “Cambridge Natural History,” “ Insects,” part ii.,
p. 293, and the original authorities cited on p. 294.

122 INTELLIGENT BEHAVIOUR

one, though, when disclosed, it almost immediately sets to
work to make funnels on the complex and perfect system”
characteristic of the species. This is but one example of
instinctive precision out of the many which could be cited.
We may say, then, that though, when an act is otherwise
shown to be intelligent, the precision is a criterion of the
level attained by the intelligence, still it cannot be said to
be a criterion which serves to distinguish intelligent from
instinctive behaviour.

Nor can we regard apparent prevision (which is sometimes
advanced as a criterion of intelligence) as specially distinctive
of intelligent acts regarded objectively in the study of animal
behaviour. For, as we have had occasion to show, there are
many instincts which display an astonishing amount of what
may be termed “blind prevision”—instance the instinctive
regard for the welfare of unborn offspring which the mother
will never see, and the instinctive preparation for an unknown
future existence in the case of insect larve.

Nor, again, is the complexity of the adjustment distinctive
of intelligence as contrasted with instinct. We have cited
examples which afford evidence of much complexity in in-
stinctive behaviour. The construction and storage of the
nest among solitary wasps, and their methods of capturing
and conveying the insects or spiders on which they prey, are
sufficiently complex. So, too, is the behaviour of the Sitaris
larva which attaches itself to the male hee, passes to the
female, and then slips on to the eggs she lays; and so, again,
is that of the Yucca moth, which collects pollen from the
anthers, conveys it to the stigma, and then lays her eggs
among the ovules. These cases show, too, that the circum-
stances may be dealt with in such a way as to extract from
them the maximum of benefit. It would be difficult intelli-
gently to improve upon the manner of dealing with the.circum-
stances displayed in many familiar modes of instinctive procedure.

There remain the novelty of the adjustment and the in-
dividuality displayed. And here we seem to have valid
criteria of intelligent behaviour. The ability to perform

INTELLIGENT BEHAVIOUR IN INSECTS 123

acts in special adaptation to new circumstances, and the
individuality manifested in dealing with the complex condi-
tions of a variable environment,—thege seem to be distinctive
features of intelligence. On the other hand, in instinctive
behaviour there seems to be no choice ; the animal is impelled
to their stereotyped performance through impulse, as by a stern
necessity ; they are so far from novel that they are performed
by every like individual of the species, and have been so per-
formed by their ancestors for generations ; and in performing
the instinctive act, the animal seems to have no more in-
dividuality or originality than a piece of adequately wound
clockwork.

Granting, then, that behaviour is shown to be intelligent by
the fact that there is evidence of profiting by experience, we
may say that the level attained by the intelligence is indicated
by the complexity of the adjustment, its precision, the in-
dividuality shown, the amount of prevision disclosed, and
in its being such as to extract from the circumstances the
maximum of benefit. Many of these points, however, serve
equally well to mark the level of instinctive procedure.

II.—InteLuicgent Benaviour in Insxcts

It is, as we have seen, among the higher invertebrates,
especially in insects, that some of the most remarkable and com-
plex instincts may be found. ‘There is,* however, a tendency
to ascribe the behaviour of insects entirely to instinct, without
sufficient evidence that neither imitation, instruction, nor
intelligent learning play any part. This is, perhaps, a survival
of the old-fashioned view that all the acts of the lower animals
are performed from ‘instinct, whereas those of human beings
are to be regarded as rational or intelligent. In popular
writings and lectures, for example, we frequently find some or
all of the following activities of ant-life ascribed to instinct :

* This and the three succeeding paragraphs are taken from “ Animal
Life and Intelligence,” p. 425:

124 INTELLIGENT BEHAVIOUR

recognition of members of the same nest ; powers of com-
munication; keeping aphides for the sake of their sweet
secretion ; collection of aphid eggs in October, hatching them
out in the nest, and taking them in the spring to the daisies,
on which they feed, for pasture; slave-making and slave-
keeping, which, in some cases, is so ancient a habit that the
enslavers are unable even to feed themselves ; keeping insects
as beasts of burden, ¢7.a kind of plant-bug to carry leaves ;
keeping beetles, etc., as domestic pets; habits of personal
cleanliness, one ant giving another a brush-up, and being
brushed-up in return; habits of play and recreation ; habits
of burying the dead ; the storage of grain and nipping the
budding rootlet to prevent further germination; the habits
described by Dr. Lincecum, and to a large extent confirmed
by Dr. McCook, that Texan ants prepare a clearing around
their nest, and six months later harvest the ant-rice, a kind of
grass of which they are particularly fond, even, according to
Lincecum, seeking and sowing the grain which shall yield this
harvest ; the collection by other ants of grass to manure the
soil on which there subsequently grows a species of fungus
upon which they feed ; the military organization of the ecitons
of Central America ; and so forth. Now, the description of
the habits of ants forms one of the most interesting chapters
in natural history. But to class them all as illustrations of
instinct is a survival of an old-fashioned method of treatment.

To put the matter in another way. Suppose that an intelli-
gent ant were to make observations on human behaviour as
displayed in one of our great cities or in an agricultural
district. Seeing so great an amount of routine work going on
around him, might he not be in danger of regarding all this
as evidence of hereditary instinct? Might he not find it
difficult to obtain satisfactory evidence of the establishment of
our habits, of the fact that this routine work has to some
extent to be learnt? Might he not say (perhaps not wholly
without truth), “I can see nothing whatever in the training
of the children of these men to fit them for their life-work.
The training of their children has no more apparent bearing

INTELLIGENT BEHAVIOUR IN INSECTS 128

upon the activities of their after-life than the feeding of our
grubs has on the duties of ant-life. And although we must
remember,” he might continue, “that these large animals do
not have the advantage which we possess of awaking suddenly,
as by a new birth, to their full faculties, still, as they grow
older, now one and now another of their deferred instincts is
unfolded and manifested. They fall into the routine of life
with little or no training as the period proper to the various
instincts arrives. If learning thereof there be, it has at
present escaped our observation. And such intelligence as
their activities evince (and many of them do show remarkable
adaptation to uniform conditions of life) would seem to be
rather ancestral than of the present time ; as is shown by the
fact that many of the adaptations are directed rather to past
conditions of life than to those which now hold good. In the
presence of new emergencies to which their instincts have not
fitted them, these poor men are often completely at a loss.
We cannot but conclude, therefore, that, although shown under
somewhat different and less favourable conditions, instinct
occupies fully as large a space in the psychology of man as it
does in that of the ant, while their intelligence is far less
unerring and, therefore, markedly inferior to our own.”

Of course, the views here attributed to the ant are very
absurd. But are they much more absurd than the views of
those who, on the evidence which we at present possess,
attribute all the varied activities of ant-life to instinct ? Take
the case of the ecitons, or military ants, or, the harvesting ants,
or the ants that are said to keep draught-bugs as beasts of
burden : have we sufficient evidence to enable us to affirm
that these modes of behaviour are purely instinctive and not
intelligent ; that all the varied manceuvres of the military
ants, for example, are displayed to the full without any learn-
ing or imitation, without teaching and without intelligence on
the part of every individual in the army.

That in some cases there is something very like a training
or education of the ant when it emerges from the pupa con-
dition is rendered probable by the observations of M. Forel.

126 INTELLIGENT BEHAVIOUR

As Romanes says,* “The young ant does not appear to come
into the world with a full instinctive knowledge of all its
duties as a member of a social community. It is led about the
nest and ‘ trained to a knowledge of domestic duties, especially
in the case of larva.’ Later on, the young ants are taught to
distinguish between friends and foes.”

We have only to weigh the evidence brought forward by
such observers as Fabre and Dr. Peckham to see that among
the solitary wasps and mason bees the behaviour, though
founded on instinct, is in large degree modified by intelligence.
The care with which a site for the tunnelled nest in the
ground is selected, betokens something more than instinct.
The following is a slightly condensed statement of Dr. and Mrs.
Peckham’s observations on one of the solitary wasps (Aporus
fasciatus).— “We were working one day in the melon-field
when we saw one of these little wasps going backwards and
dragging a spider. She twice left it on the ground while she
circled about for a moment, but soon carried it up on to one
of the large melon leaves, and left it there while she made a
long and careful study of the locality, skimming close to the
ground in and out among the vines; at length she went under
a leaf close to the ground, and began to dig. After her head
was well down in the ground we broke off the leaf that we
might see her method of work. She went on for ten minutes
without noticing the change, and then, without any circling,
flew off to visit her spider. When she tried to return to her
hole it was evident that some landmark was missing, Again
and again she zig-zagged from the spider to the nesting-place,
going by a sort of path among the vines from leaf to leaf, and
from blossom to blossom, but when she reached ‘the spot she
did not recognize it. At last we laid the leaf back in its place
over the opening, when she at once went in and resumed her
work, keeping at it steadily for ten minutes longer. At this
point she suddenly reversed her operations, and began to fill in
the hole that she had made. She then glanced at the spider,

* « Animal Intelligence,” p. 59.
+ “Instincts and Habits of the Solitary Wasps,” p. 55.

INTELLIGENT BEHAVIOUR IN INSECTS 127

selected a new place, and began to dig again. This hole was
also filled in ; she looked once more at the spider, and started
a nest in a new place. This, in turn, was soon abandoned, as
was a fourth. The fifth beginning was made under a leaf
that lay close to the ground, but after twenty minutes’ work
this place also was abandoned and a sixth started. This, how-
ever, was the final choice, and after forty-five minutes spent in
digging it was completed.”

This description shows an amount of apparent fastidious-
ness which is quite irreconcilable with the hypothesis that the
behaviour is merely instinctive. Not less fastidious are some
wasps in the temporary closure of the hole with a stone or
pellet of earth, the operation being repeated several times

Fic. 19.—Solitary Wasp using a stone to beat down the earth
over its nest (after Peckham).

with different covers before the insect seems to be satisfied ;
while in other cases the hole is hidden by bringing earth
in such quantity as to render the place indistinguishable
from the rest of the field. But in one case observed. by Dr.
Peckham, intelligent procedure was carried so far as ap-
parently to involve the use of a tool, the same behaviour
having been independently observed in the same genus (Am-
mophila) by Dr. 8. W. Williston of Kansas University. ‘“ Just

128 INTELLIGENT BEHAVIOUR

here,” writes Dr. Peckham,* “must be told the story of one
little wasp whose individuality stands out in our minds more
distinctly than that of any of the others. In filling up her
nest she put her head down into it and bit away the loose
earth from the sides, letting it fall to the bottom of the
burrow, and then, after a quantity had accumulated, jammed
it down with her head. Earth was then brought from the
outside and pressed in, and then more was bitten from the
sides. When, at last, the filling was level with the ground,
she brought a quantity of fine grains of dirt to the spot, and,
picking up a small pebble in her mandibles, used it as a
hammer in pounding them down with rapid strokes, thus
making this spot as hard and firm as the surrounding surface.
Before we could recover from our astonishment at this per-
formance she had dropped her stone and was bringing more
earth, and in a moment we saw her pick up the pebble and
again pound the earth into place with it. Once more the
whole process was repeated, and then the little creature flew
away.”

Here we have intelligent behaviour rising to a level to
which some would apply the term rational. For the act may
be held to afford evidence of the perception of the relation
of the means employed to an end to be attained, and some
general conception of purpose. In this section, which deals
with description of behaviour based on observation, the
psychological explanation cannot be discussed. Similar in-
dications of deliberate action may be held to be afforded by
the sometimes elaborate “ locality studies” which these insects
seem to make,—by the “ care that is taken by wasps to acquaint
themselves with the surroundings of their nests.” A Sphez,
for example, which had partially made and then abandoned
several nests, left them without any locality study ; but when
she had completed a nest in a suitable spot she made “a most
thorough and systematic study of the surroundings. She flew
in and out among the plants, first in narrow circles near the
surface of the ground, and then in wider and wider ones as

* Op, cit., p. 22,

INTELLIGENT BEHAVIOUR IN INSECTS 129

she rose higher in the air, until at last she took a straight line
and disappeared in the distance.” Another species (Cerceris
deserta) “has the habit of making a number of half circles in
front of the nest, and then, after rising a little higher, of
flying several times completely round it.” The method of
procedure is, it seems, so normal to the species that it is pro-
bably founded on an instinctive basis. Dr. and Mrs, Peckham,
in commenting on their observations, say : * “If the examination
of the objects about the nest makes no impression upon the
wasp, or if it is not remembered, she ought not to be incon-
venienced nor thrown off her track when weeds and stones
are removed and the surface of the ground is smoothed over ;
but this is just what happens.” For convenience of observa-
tion they “sometimes gently moved intercepting objects to
one side, but even such a slight change threw the wasp out
of her bearings, and made it difficult for her to recover her
treasure.” Where wasps form a number of nests in a small
plot of ground, as in the case of Bember, each knows and
returns to its own hole, as was proved by Dr. Peckham, who
marked the insects and their nests with paint.

So, too, with regard to prey. In the course of his observa-
tions on Pompilus, Fabre removed the spider which the wasp
had deposited on a tuft of vegetation before she made her
nest. As she was at work beneath the surface she could not
see what went on above ground or where the spider had been
redeposited some twenty inches from its former position. On
emerging from the nest the wasp went straight to the original
spot, searched round it for some time, then made further
excursions, and discovered the spider. After slightly altering
its position, and placing it on another tuft of vegetation, she
returned to her subterranean labours, giving the observer an
opportunity of again moving the spider. Five times did
Fabre repeat the operation, and every time the wasp returned
to the spot where she had last deposited her prey.

The same observer records some interesting experiments
with the mason bee, Chalicodoma. The mud nests of the species

* Op. ctt., p. 215.
K

130 INTELLIGENT BEHAVIOUR

investigated were affixed to stones on the banks of the Rhone.
When a nest was partially constructed, the bee having flown
off for more material, Fabre moved the stone to a new position,
near at hand and easily visible from the original site. The
bee went straight to the place where the nest had been,
searched the immediate neighbourhood, flew off, and returned
to the same spot to continue the search. If she came across
her own nest in its new position she did not recognize it as
hers, but left it after examination. But if a stone with the
nest of another bee in about the same stage of construction
was placed in the position occupied by her own, she adopted
it. And when two nests near together, both half built, were
transposed, each bee unhesitatingly adopted the nest which
occupied the position where its own nest had been. It may
well seem strange that, the general locality-memory being so
well marked, the recognition of the particular stone and nest
should be deficient. This may be due to the fact that the
so-called compound eyes are the organs concerned in locality
vision, while the ocelli deal with details at very close range,
and that the former alone afford the requisite data for recog-
nition; by their instrumentality alone arises the conscious
situation which affords guidance in behaviour. And in that
situation slight changes which for us make it “still the same
but with a difference” render it no longer the same for a
being of more limited intelligence—one probably incapable
of analyzing the situation and seeing that the sameness
preponderates over the difference. Be this as it may, the
failure of a bee to recognize its own nest under circumstances
so foreign to its experience as removal to a new spot may be
paralleled with what I have observed in the case of sticklebacks.
A nest had been built in a round glass bell jar which stood
near a window. Some aquatic vegetation grew in the tank,
and the nest was built on the window side. An experiment was
made by turning the large bell jar through a right angle. The
male stickleback searched for its nest in the old direction on
the window side—that is to say, the same position in reference

INTELLIGENT BEHAVIOUR IN INSECTS 131

to the incidence of the light. The search was, of course,
fruitless, and a new nest was begun in this position. Presently
the old nest was discovered, and was then vigorously destroyed
in just the same way as the nest of a rival is pulled to pieces
and scattered. Here a new incidence of light and new direction
of shadows seemed to have completely transformed the visual
situation.

To return to insects, it is probable that the homing faculty
is not the result of an inborn mysterious instinct dependent
on some sense of direction of which we have no knowledge,
but is based upon experience gained during their flight hither
and thither—that, in a word, it is intelligent and not instinc-
tive. Experiments of Fabre at first seemed to suggest some
magnetic influence to which bees were sensitive; for when a
minute magnet was fixed to a bee as it started on its return
journey, the insect was at fault; but as a check experiment
he affixed a piece of straw instead of a magnet, with similar
results. Some of Fabre’s observations and those of Dr.
Bethe* are difficult to reconcile with the hypothesis that, in
the homing, guidance is due to acquired acquaintance with
the locality. But, on the other hand, the experiments of
Lord Avebury and of Romanes seem to favour this view.
Romanes found that when bees were taken inland from their
hive near the seaboard, and then liberated, they returned
from considerable distances, the whole locality being familiar ;
but taken to the seashore, where the objects around them
were unfamiliar (since the seashore is not the place where
flowers and nectar are to be found), the bees, though not far
distant from the hive, were nonplussed and lost their way.
Dr. and Mrs. Peckham, as the result of their extremely careful
observations, seem to have no doubt that the homing of solitary
wasps is due to locality-experience ; and of the social wasp,
Polistes fusca, they say:t ‘“ We have seen the young workers
make repeated locality studies when they first began to venture

* “Diirfen wir den Ameisen und Bienen psychische Qualitaten

Zuschreiben.” Pfluger’s Archiv., 1xx., 1898.
t “Solitary Wasps,” p. 219.

132 INTELLIGENT BEHAVIOUR

away from home, but as they occupy the same nest all summer
they, of course, grow more and more familiar with their sur-
roundings, until they become so thoroughly acquainted with
them that they can find their way without the least difficulty.
We have no doubt that with them, as with the solitary wasps,
the faculty is not instinctive, but is the direct outcome of
individual experience.”

In the interesting pages of the works in-which Dr. and Mrs.
Peckham describe their investigations, there are many observa-
tions which show that wasps are capable of intelligently profiting
by the experience which their instinctive behaviour places them
ina position toacquire. The inherited tendencies and aptitudes
pave the way for acquired modification and accommodation of
behaviour. To catch and paralyze spiders, to dig and prepare
a tunnelled nest, and to carry the prey to the nest, all this
affords the instinctive basis ; but when the observers tell us
that they ‘“‘ have several times seen wasps enlarge their holes
when a trial had demonstrated that a spider would not go in,”
and even on one occasion without trial when an unusually
bulky spider was brought, there is something beyond instinct ;
there is intelligent adjustment to special circumstances given
in experience. Presumably intelligent is the habit frequently
observed in one species of Pompilus, and occasionally in another,
of hanging the paralyzed spider in a crotch of a branching
stem, usually of bean or sorrel, where it will be safe from the
depredation of ants. On one occasion Dr. Peckham, desirous
of seeing the exact manner in which the victim was stung,
substituted an unhurt spider for that which the wasp had
paralyzed.* “According to the habit of its species when
danger threatens, it kept perfectly quiet, and when the wasp
returned it was hanging there as motionless as a piece of dead
matter ; but she would not touch it; she hunted all over that
plant and then over several others near to it, returning con-
tinually to look again at the right spot. After five minutes
she flew off in the direction of the woods to catch another
spider. Why did she go to the woods? Why did she not

* Op. cit., pp. 131, 132.

INTELLIGENT BEHAVIOUR IN INSECTS 133

take the one that hung there in plain view? It could not
have been due to the fact that we had handled the spider,
since when, on other occasions, we took one that had been
paralyzed, examined it, and then returned it to the wasp, she
accepted it without hesitation. . . . In forty minutes she came
back with another spider, but, instead of taking it into the nest,

Fic. 20.—Spiders placed by Solitary Wasps in the crotches of
branching stems (after Peckham).

she hung it upon a bean plant near by, and then proceeded to
dig a new hole a few inches distant from the first. Foolish
little wasp, what a waste of labour! . Truly, if you are endowed
with energy beyond your fellows, you are but meagrely furnished
with reason.”

Here we have the routine of instinct—the normal mode of
hunting and capturing prey, the normal procedure of bringing
the spider, and then making the nest, predominating over any
tendency to initiate intelligent improvements. This, however,
should not surprise us, in whom the force of habit is often so
strong. Nor should we feel surprise at the apparently stupid
tolerance some of these wasps display in presence of parasites.

134 INTELLIGENT BEHAVIOUR

Bembex, which does not store and close its cell, but brings
continual supplies of food to its larvae, is not disturbed by the
presence in the cell of the grubs of the parasitic fly Multo-
gramma. She could, we think, easily free her nest of these
intruders, but she continues to bring supplies, though the
parasites may absorb it all and leave her own Jlarvee to perish.
She adapts her procedure to the new conditions, being in-
capable of knowing that she is feeding the enemies of her
race.

Enough has now been said to show the extent and the
limitations of the intelligence of such insects as the solitary
wasps. It will be noticed that the acquired modifications of
behaviour occur in close connection with the inherited ground-
plan of instinctive procedure. We shall have occasion to note
the same connection in our discussion of social behaviour in
the next chapter. And we shall consider the influence of
intelligence on instinct before we bring this chapter to a
close. ;

III.—Some Resuuts or EXPERIMENT

Tt is unnecessary to give a resumé of entertaining anecdotes
illustrative of intelligent behaviour in the higher animals.
Such anecdotes are too often the outcome of casual observa-
tions by untrained observers ; and the interpretation put upon
the facts frequently shows a want of psychological discrimina-
tion. Such is not the material of which science is constituted.
What is needed is systematic observation conducted, so far as
possible, under controlled conditions. Two things are neces-
sary : first, to distinguish instinctive behaviour, inherited as
such, from the acquired modifications or new departures due
to intelligence ; and secondly, to determine the method and
range of intelligent procedure. These problems can only be
solved in their entirety by a complete knowledge of the life-
history of the animal concerned. But they may be attacked
in detail by a systematic study of particular modes of behaviour,
and by an investigation into their manner of origin. That

SOME RESULTS OF EXPERIMENT 135

this may be done with some approach to accuracy, resort must
be had to experiment, which permits of observation under
controlled conditions.

To ascertain, for example, how far nest-building is instinc-
tive in birds, Mr. John 8. Budgett hatched a hen greenfinch
under a canary. In the following autumn he bought a caged
bird, a cock, probably of the same year, and in the succeeding
spring turned the pair into a large aviary, supplying such
material as twigs, rootlets, dried grass, moss, feathers, sheep’s
wool and horsehair. The hen soon began to build, the cock
bird taking no share in the work, and finished her nest in a
few days. On careful comparison it was found to resemble
that of a wild greenfinch in every particular, being made of
wool, roots, and moss, lined with horsehair. A second nest the
aviary greenfinch built was also quite normal.

In the case of a bullfinch which Mr. Budgett reared, having
obtained it when a few weeks’ old, the first nest was composed
of dried grass with a little wool and hair, but without either
rootlets or twigs. A second which she built was, however,
quite typical, made of fine twigs and roots, and lined with
horsehair ; as was also a third nest.

It is just possible, though most improbable, that the bull-
finch utilized its three weeks’ experience gained in the nest
from which it was taken. But Mr. Jenner Weir describes *
observations on canaries in which this source of experience is
excluded. “It is usual,” he says, “ with canary fanciers to
take out the nest constructed by, the parent birds, and to place
a felt. nest in its place, and when her young are hatched, and
old enough to be handled, to place a second clean nest, also of
felt, in the box, removing the other. But I never knew that
canaries so reared failed to make a nest when the breeding
time arrived. I have, on the other hand, marvelled to see
how like a wild bird’s the nests are constructed. It is custo-
mary to supply them with a small set of materials, such as
moss and hair. They use the moss for the foundation, and

* Ina letter to Darwin, quoted by Romanes, “Mental Evolution in
Animals,” p. 226,

136 INTELLIGENT BEHAVIOUR

line with the finer materials, just as a wild goldfinch would
do.”

Experiment seems, therefore, to show in a way, and with a
clearness impossible of attainment by observation under natural
uncontrolled conditions, that nest-building in birds is instinc-
tive. That the manner and method of procedure is often
modified in accordance with special conditions—that the in-
stinctive outline of nidification receives its final touches
through individual experience—is sometimes seen under
nature, and more often under the semi-experimental conditions
of domestication. Thus three pairs of pigeons in the Wilson
Tower of Clifton College made their nests in 1898, as I am
informed by Mr. H. C. Playne, of galvanized iron wire, pieces
of which were left in a corner at the top of the tower, thus
affording a parallel to the behaviour of the unconventional
crow of Calcutta, mentioned by Mr. J. W. Headley,* which
made its nest of soda-water bottle wires, which it picked up in
a back yard. But even in this matter experiment serves to
bring out clearly the selective influence which is exercised by
intelligence. Bolton,f in 1792, observed a pair of goldfinches
beginning to build their nest in his garden. They formed the
ground-work of moss, grass, etc., as usual ; but on his scatter-
ing small parcels of wool in different parts of the garden they,
in great measure, left off the use of their own stuff and used
the wool. Afterwards he gave them cotton, and they then
used this instead of the wool; then he supplied fine down, and
they finished their work with this, leaving the wool and
cotton.

Tn studying the behaviour of wild animals under natural
conditions, it must always be difficult to distinguish the con-
genital basis from the acquired elements ; for both tend to
bring about a working adjustment to the conditions of life,
and we can seldom have opportunities of tracing the, interplay
of the factors which produce the instinct-habits of adult life.

* « The Structure and Life of Birds,” p. 335.
+ Preface to “ Harmonia Ruralis,” quoted by Yarrell, “ British Birds,”
vol. i., p. 541, ‘

SOME RESULTS OF EXPERIMENT 137

But under domestication we seek to bring about a new working
adjustment to conditions imposed by man. The skilful trainer
utilizes the natural instinctive tendencies as a basis ; and, by
a system of rewards and punishments, leads the intelligent
modifications of behaviour along lines directed by his deliberate
purpose. The conditions are largely those of experiment,
and they bring out the play and range of intelligence in a way
which would otherwise elude our observation. The training
of falcons for the chase affords a good illustration, since they
cannot be bred in confinement, and the effects of training
cannot therefore be hereditary. The falconer’s object is to
modify the congenital instinctive behaviour of a bird of prey
for the purposes of sport. She is trained to the lure at first
at short distances, and step by step through longer flights ; she
is taught by snatching away the lure to stoop at it repeatedly
as often as it is jerked aside ; and then she is trained on living
quarry, at first under easy conditions, till eventually she can be
flown ata wild bird. And as a result a well-trained falcon will
follow her master from field to field, regulating her flight by
his movements, always ready for a stoop when the quarry is
sprung. The fact that she can be thus educated for her work
shows that her behaviour is plastic, and can be moulded by
intelligence. Experimental conditions reveal the fact ; but
under nature the moulding influence of intelligence is pre-
sumably not less important, though it is more directly in line
with the congenital instinctive tendencies.

That much of the behaviour of the higher animals is guided
by experience similar to that which plays so large a part in
their training under the experimental conditions of domestica-
tion is generally admitted. But what are the range and
limits of animal intelligence, and whether it attains the level
of rational conduct, in the restricted sense of the term
“yational,” are questions open to discussion, to which answers
are more likely to be obtained through experiment than by
chance observations.

Before giving some of the results of such experiment it
will be well to revert to the distinction, which was drawn in the

138 INTELLIGENT BEHAVIOUR

second chapter, between the lower or intelligent stage of mental
development and the higher or rational stage. It will be
remembered that rational processes were characterized by the
fact that the situations contain the products of reflective
thought, presumably absent in the earlier stages of develop-
ment ; that they were further characterized by a new purpose
or end of consciousness, namely, to explain the situations which
at an earlier stage are merely accepted as they are given in
presentation or re-presentation ; they require deliberate atten-
tion to the relationships which hold good among the several
elements of successive situations ; and they involve, so far as
behaviour is concerned, the intentional application of an ideal
scheme with the object of rational guidance.

On the other hand, the animal at the stage of intelligent
behaviour deals with the circumstances of his comparatively
simple life by making use of the particular situations which
have been presented to consciousness in the course of his prac-
tical experience. If such an animal be placed in the midst of
new circumstances he has to find out by a process of trial and
error how they are to be met. After a longer or shorter
period of trial, guided only by particular experiences, he
chances to hit upon a mode of procedure which is successful.
The successful act is then incorporated in a new situation ; at
first, perhaps, only incompletely.. The association is eventually
established by repetition, through which is acquired the habit
of doing the right thing in the appropriate manner. Why he
does this and not something else, in so far as he is intelligent
and not rational, he probably neither knows nor has the wit
to consider. The satisfaction of success suffices for intelligence
as such. If the circumstances be so modified as to render the
particular mode of meeting them ineffectual, after trying
again and again in the old way, he will sometimes stumble
upon the proper mode of overcoming the difficulty, and after
doing so two or three times a new conscious situation involving
the requisite associations will be established, and the appro-
priate behaviour will become habitual. But why this new
mode of procedure rather than any other is adopted, intelligence

SOME RESULTS OF EXPERIMENT 139

as such does not know, because it does not analyze the
situation and disentangle the essential relationships. The
satisfaction of success again suffices. In a word, such an
animal in the perceptual stage of mental development seems
wanting in the power of reflection. He does not appear to show
evidence of framing anything like a general scheme of know-
ledge which he can apply to the solution of particular problems,
of a practical nature, involving difficulties and obstacles.

The method of intelligence—in the sense in which we are
using this term—the method of varied trial and error with the
utilization of chance success, ig a lengthy and somewhat
clumsy process; but it suffices. Now contrast it with the
procedure of a rational animal, such as man is or may. be.
When he is confronted by a difficulty he is not content to meet
it by trying this way, and that way, and another way, anyhow,
and trusting to chance to bring success, but he considers the
problem in all its relations with a view to ascertaining the
essential nature of the difficulty. For each attempted mode of
meeting the case he has a definite reason. He knows why
he does this and not that. He has a plan or scheme which he
puts into execution. And if it fail, he is not content till
he finds out wherein the failure lay. This enables him to plan
a better scheme. He sees why it is better; and if at last he
be successful by a happy hit, as in the chance procedure of
intelligence, he looks for the reason of it. And seeing why
this fortunate attempt, unlike his previous efforts, just meets
the case, he repeats it because he perceives that herein lies the
essential solution of the difficulty. Both in the case of intelli- |
gence and in that of reason, as here distinguished, present
procedure is based upon past experience; but reason has built
upon the foundations thus laid an orderly scheme, and knows
its whys and wherefores, while intelligence is at the mercy of
chance associations. The reason for success it has not the wit
to assign. ‘

The essential difference between the two cases may be put
in another way by saying that the intelligent being forms
sensory impressions and sensory images linked together by

140 INTELLIGENT BEHAVIOUR

bonds of association, combining and coalescing to constitute a
conscious situation effective in behaviour under the guiding
influence of pleasure and pain; while the rational being not
only does all this, but goes further. He fixes his attention on
the way in which the elements in the situation are con-
nected and related; he builds an ideal framework on which
the sensory impressions are set or move in an orderly manner.
And it is this scheme, fashioned by reason and transforming
the situation, which he utilizes in dealing with difficulties.

Yet another way of putting the same essential distinction
is to say that intelligence deals with pictures, either directly
presented to the senses or called up in re-presentation. If we
state the matter thus, however, we must remember that the
“pictures”? may be painted in colours supplied by any of the
senses ; and that smells, tastes, sounds, touches, pressures,
limb-movements, and so forth, are elements in the pictured
product. Bearing this in mind, we may say that intelligence
deals with sensory impressions and their revived images in
concrete and particular situations ; while reason analyzes the
pictures, and extracts from them general notions in terms of
which the pictures may be explained. For example, we picture
a stone falling to the earth; but we explain it by the general
notion of gravitative attraction. The conception forms part
of our ideal scheme of knowledge, which is not itself picturable,
though this or that example of its action may be presented or
re-presented in sensory imagery.

Once more: we may say—and this way of looking at the
question arises naturally out of what has gone before—that
intelligence deals with concrete examples, and does not rise to
the abstract and general rule. The ideal scheme of reason is
the result of abstraction and generalization. It is a frame-
work of conceptions which can be applied to the particular
facts which fall under observation to see whether it fits and
meets the case. Intelligence has to deal with the facts as they
present themselves, without the aid of an organized system of
knowledge built up into an ideal scheme. ,

Enough has now been said to indicate the distinction

SOME RESULTS OF EXPERIMENT 141

between the method of intelligence and that of reason. It
may, no doubt, be said that the terminology used is open to
criticism ; for, on the one hand, the words “ intelligence ” and
“intelligent” are frequently used as synonymous with “reason ”
and “rational ;” and, on the other hand, acts requiring neither
abstraction, generalization, nor the application of any scheme
of knowledge are frequently spoken of as “rational.” Hence
there is, it may be urged, some danger of misunderstanding.
This may be granted. And unless some such restriction of
meaning under suitable terms be accepted by psychologists,
misunderstanding will continue. More essential, however, than
the distinctive terms we are to use is the distinction of method
which underlies them. That, I trust, is sound. Dr. Lindley,
in an interesting paper on “‘ A Study of Puzzles,” * has utilized
the distinction in his investigation of the mental development
of children, and has found that the procedure of young children
is predominantly of the “‘sense-trial and error” order which
has above been termed intelligent; and he expresses the
opinion that “most of the adaptations of animals are on
this sense-trial and error level.”

Such certainly seems to be the conclusion to be drawn from
my own experimental observations on dogs. It has frequently
been asserted that the behaviour of a dog with a stick in his
mouth, when he comes to a narrow gap, shows that he at once
perceives the nature of the difficulty, and meets it in a rational
manner by adopting the appropriate plan of action. He pulls
the stick through by one end. But experiments, which I have
elsewhere described,t showed that a fox terrier, fourteen months
old, seemed to be incapable of perceiving the nature of the
difficulty which vertical iron railings presented to his passage
with a stick in his mouth, and only imperfectly learnt to over-
come it after many ineffectual trials and many failures. The
results obtained on the first afternoon may be quoted to indicate
the nature of the evidence. The dog was sent after a short stick
into a field, and had to pass through vertical rails about six

* American Journal of Psychology, vol. viii., no. 4, pp. 431-493.
+ “Introduction to Comparative Psychology,” p. 255.

142 INTELLIGENT BEHAVIOUR

inches apart. On his return the stick caught at the ends. I
whistled and turned as if to leave; and the dog pushed and
struggled vigorously. He then retired into the field, lay down,
and began gnawing the stick, but, when called, came slowly up
to the railings and stuck again. After some efforts he put his
head on one side, and brought the stick, a short one, through.
After patting and encouraging him, I sent him after it again.
On his return he came up to the railings with more confidence,
but, holding the stick by the middle, found his passage barred.
After some struggles he dropped it and came through without
it. Sent after it again, he put his head through the railings,
seized the stick by the middle, and then pulled with all his
might, dancing up and down in his endeavours to effect a
passage. Turning his head in his efforts, he at last brought
the stick through. A third time he was again foiled; again
dropped the stick ; and again seizing it by the middle tried to pull
it through. I then placed the stick so that he could easily seize
it by one end and draw it through the opening between the
rails. But when I sent him after it, he went through into the
field, picked up the stick by the middle, and tried to push his
way between the railings, succeeding, after many abortive
attempts, by holding his head on one side.

Subsequent trials on many occasions yielded similar results.
But the following summer, when I resumed the experiments, I
was able with some guidance to teach him to bring a long stick
to the railings, drop it, and then draw the stick through by one
end; though even then, if he had dropped it so that one end
just caught a rail, he often failed, shaking his head vigorously,
dropping the stick and seizing it again, and repeating this
behaviour until it chanced to fall in a more favourable position.
He did not apparently perceive that by gently moving the stick
a little one way or other the difficulty could be simply overcome
with little effort. Nor when given a crooked stick, which caught
in a rail, did he show any sign of perceiving that by pushing the
stick and freeing the crook he could pull the stick through.
Each time the crook caught he pulled with all his strength,
seizing the stick now at the end, now in the middle, and now

SOME RESULTS OF EXPERIMENT 143

near the crook. At length he seized the crook itself, and with
a wrench broke it off. A man who was passing, and who had
paused for a couple of minutes to watch the proceedings, said,
“Clever dog that, sir; he knows where the hitch do lie.” The
remark was the characteristic outcome of two minutes’ chance
observation. During the half hour or more during which I
had watched the dog he had tried nearly every possible way of
holding and tugging at the stick. Such is the mode of
behaviour based on intelligence—continued trial and failure,
until a happy effect is reached, not by methodically planning,
but by chance.

Two of my friends criticized these results, and said that
they only showed how stupid my dog was. Their dogs would
have acted very differently. I suggested that the question
could easily be put to the test of experiment. The behaviour
of the dog was in each cage—the one a very intelligent York-
shire terrier, the other an English terrier—similar to that
above described. The owner of the latter was somewhat

. annoyed, used forcible language, and told the dog that he
could do it perfectly well if he tried.

In experimenting with my fox terrier on the method
adopted in seizing and carrying differently balanced objects,
I used (1) a straight stick, the centre of gravity of which was
at the middle; (2) a Kaffir knob-kerrie, the centre of gravity
of which was about six inches from the knob ; (3) a light
geological hammer; and (4) a heavier hammer. In the last,
the centre of balance was close to the hammer head. The net
result of the observations was that the best place for seizing
and holding the object was hit upon in each case after indefinite
trials ; that after three or four days’ continuous experience with
one (say the knob-kerrie), another (say the stick) was at first
seized nearer one end, showing the influence of the more recent
association ; and that there was little indication of the dog’s
seizing any one of the four at once in the right place, that is
to say, the point of seizure was not clearly differentiated in
accordance with the look of the object. I tied a piece of
string, in later trials, round the centre of balance, but this, at

144 INTELLIGENT BEHAVIOUR

the time of the dog’s death, had not served as a sure guide to
his experience.

The way in which my dog learnt to lift the latch of the
garden gate, and thus let himself out, affords a good example
of intelligent behaviour. The iron gate outside my house is
held to by a latch, but swings open by its own weight if the
latch be lifted. Whenever he wanted to go ont the fox
terrier raised the latch with the back of his head, and thus
released the gate, which swung open. Now the question
in any such case is: How did he learn the trick? In this
particular case the question can be answered, because he was
carefully watched. When he was put outside the door, he
naturally wanted to get out into the road, where there was
much to tempt him—the chance of a run, other dogs to sniff
at, possibly cats to be worried. He gazed eagerly out through
the railings on the low parapet wall shown in the illustration ;
and in due time chanced to gaze out under the latch, lifting
it with his head. He withdrew his head and looked out
elsewhere ; but the gate had swung open. Here was the
fortunate occurrence arising out of natural tendencies in a dog.
But the association between looking out just there and the
open gate with a free passage into the road is somewhat
indirect. The coalescence of the presentative and re-pre-
sentative elements into a conscious situation effective for the
guidance of behaviour was not effected at once. After some
ten or twelve experiences, in each of which the exit was more
rapidly effected with less gazing out at wrong places, the fox
terrier had learnt to go straight and without hesitation to
the right spot. In this case the lifting of the latch was
unquestionably hit on by accident, and the trick was only
rendered habitual by repeated association in the same situation
of the chance act and the happy escape. Once firmly
established, however, the behaviour remained constant through-
out the remainder of the dog’s life, some five or six years.

Mr. E. J. Shellard observed * an act of similar import in
a Scotch staghound, which “appeared at first to be the result

* “Tntroduction to Comparative Psychology,” p. 290.

SOME RESULTS OF EXPERIMENT 145

of thought,” but which, on closer observation, was clearly seen
to be the result of intelligence in the restricted sense of the

¥ 1c, 21.—Fox-terrier lifting the latch of a gate.

term. The dog released the lever-latch of a yard door. “ At
first he raised his paws to the door and scratched violently,

manifesting various signs of impatience. His scratches, which
L

146 INTELLIGENT BEHAVIOUR

extended from the top of the door downwards, and over the
whole area, would thus inevitably at some time or other reach the
handle of the latch, which was thus struck forcibly downwards,
the latch itself rising upwards. The door would then open
from the weight of the dog pushing against it. The dog
always opened the door in this manner from the time when
the incident was first noticed until he left, a period of about
three years. The door was opened with no greater ease at
the expiration of that period than at the commencement.
His paws would strike various parts of the door, and he never
appeared to exercise any degree of judgment in the localization
of his strokes, the fact of his paws striking the handle of the
latch being a necessary result, providing the dog had sufficient
patience and strength to continue.”

One or two more experiments with my fox terrier may be
briefly described. I watched his behaviour when a solid
indiarubber ball was thrown towards a wall standing at right
angles to its course. At first he followed it right up to the
‘wall and then back as it rebounded. So long as it travelled
with such velocity as to be only just ahead of him he pursued
the same course. But when it was thrown more violently,
so as to meet him on the rebound as he ran towards the wall,
he learnt that he was thus able to seize it as it came
towards him. And, profiting by the incidental experience
thus gained, he acquired the habit—though for long with
some uncertainty of reaction—of slowing off when the
object of his pursuit reached the wall so as to await its
rebound. Again, when the ball was thrown so as to glance
at a wide angle from a surface, at first—when the velocity
was such as to keep it just ahead of him—he followed its
course. But when the velocity was increased he learnt to
take a short cut along the third side of a triangle, so as to
catch the object at some distance from the wall. A third
series of experiments were made where a right angle was
formed by the meeting of two surfaces. One side of the
angle, the left, was dealt with for a day or two. At first the
ball was directly followed. Then a short cut was taken to

SOME RESULTS OF EXPERIMENT 147

meet its deflected course. On the fourth day this method was
well established. On the fifth, the ball was thrown so as to
strike the other or right side of the angle, and thus be deflected
in the opposite direction. The dog followed the old course
(the short cut to the left) and was completely non-plussed,
searching that side, then more widely, and not finding the
ball for eleven minutes. On repeating the experiment thrice,
similar results were that day obtained. On the following day
the ball was thrown just ahead of him, so as to strike to the
right of the angle, and was followed and caught. This course
was pursued for three days, and he then learnt to take a short
cut to the right. On the next day the ball was sent, as at
first, to the left, and the dog was again non-plussed. I did
not succeed in getting him to associate a given difference of
initial direction with a resultant difference of deflection.

I may here mention that, whenever searching for a ball of
which he had lost sight in the road, he would run along the
gutter first on one side and then on the other. A friend who
was walking with me one day regarded this as a clear case of
rational inference. ‘The dog knows,” he said, “the effects
of the convex curvature of the road as well as we do.” I am
convinced, however (having watched his ways from a puppy),
that this method of search was gradually established on a
basis of practical experience. No logical inference on his
part is necessary for the interpretation of the facts; and we
should not assume its presence unless the evidence compels us
to do so.

Dr. E. L. Thorndike, in a monograph on “ Animal Intelli-
gence” published as a supplement to the Psychological Review
(June, 1898), has -fully described and carefully discussed a
number of interesting experiments. The subjects (one might,
alas! almost say victims) of some of these were thirteen
kittens or cats from three to eighteen months old. His
method of investigation shall be stated in his own words.

“ After considerable preliminary observation of animals’ behaviour
under various conditions, I chose for my general method one which,

148 INTELLIGENT BEHAVIOUR

simple as it is, possesses several other marked advantages besides
those which accompany experiment of any sort. It was merely to
put animals when hungry in enclosures from which they could escape
by some simple act, such as pulling at a loop of cord, pressing a lever,
or stepping on a platform. The animal was put in the enclosure, food
was left outside in sight, and his actions observed. Besides recording
his general behaviour, special notice was taken of how he succeeded
in doing the necessary act (in case he did succeed), and a record
was kept of the time that he was.in the box before performing the
successful pull, or clawing, or bite. This was repeated until the
animal] had formed a perfect association between the sense-impression
of the interior of that box and the impulse leading to the successful
movement. When the association was thus perfect, the time taken
to escape was, of course, practically constant and very short.

“_ J ft jd |

IIH _JFS
Fic, 22.—Cage used in Professor Thorndike’s experiments.

)

“Tf, on the other hand, after a certain time the animal did not
succeed, he was taken out, but not fed. If, after a sufficient number
of trials, he failed to get out, the case was recorded as one of complete
failure. Enough different sorts of methods of escape were tried to
make it fairly sure that association in general, not association of a
particular sort of impulse, was being studied. Enough animals were
taken with each box or pen to make it sure that the results were not
due to individual peculiarities. None of the animals used had any
previous acquaintance with any of the mechanical contrivances by
which the doors were opened. So far as possible the animals were
kept in a uniform state of hunger, which was practically utter hunger.”

SOME RESULTS OF EXPERIMENT 149

To Dr. Thorndike’s monograph we must refer those who
desire detailed information as to apparatus and procedure. It
must here suffice to state that the box-cages employed were
rudely constructed of wooden laths, and formed cramped
prisons about twenty inches long by fifteen broad and twelve
high. Nine contained such simple mechanisms as Dr. Thorn-
dike describes in the passage above quoted. When a loop or
cord was pulled, a button turned, or a lever depressed, the
door fell open. In another, pressure on the door as well as
depression of a thumb-latch was required. In one cage two
simple acts on the part of the kitten were necessary, pulling a
cord and pushing aside a piece of board; and in yet others
three acts were requisite. In those boxes from which escape
was more difficult a few of the cats failed to get out. The
times occupied in thoroughly learning the trick of the box by
those who were successful are plotted in a series of curves,
the essential feature of which is the graphic expression of a
gradual diminution in the time interval between imprisonment
and escape in successive trials. This is shown in Fig. 23,
which is constructed from some of Dr. Thorndike’s data. In
some cases the cats were set free from a box when they (1)
licked themselves or (2) scratched themselves.

Dr. Thorndike comments on the results of his experiments
as follows :—

“When put into the box the cat would show evident signs of dis-
comfort and of an impulse to escape from confinement. It tries to
squeeze through any opening; it claws and bites at the bars or wire;
it thrusts its paws out through any opening, and claws at everything it
reaches; it continues its efforts when it strikes anything loose and
shaky: it may claw at things within the box. It does not pay very
much attention to the food outside, but seems simply to strive in-
stinctively to escape from confinement. ‘The vigour with which it
struggles is extraordinary. For eight or ten minutes it will claw, and
bite, and squeeze incessantly. . . The cat that is clawing all over the
box in her impulsive struggle will probably claw the string, or loop, or
button so as to open the door. And gradually all the other non-
successful impulses will be stamped out, and the particular impulse
leading to the successful act will be stamped in by the resulting

150 INTELLIGENT BEHAVIOUR

pleasure, until, after many trials, the cat will, when put in the box,
‘immediately claw the button or loop in a definite way. . . . Starting,
then, with its store of instinctive impulses, the cat hits upon the

Scale of Time in Seconds

Tt T 7 T T T T

-— & oe ee Sy am

a3 § € 8 $< ss ss FS sRERESS
« | 7160
Si
s | 30
Bg
S$ | 90
«
mT 60
s
oe OQe——q—
ae Be
ag
= T 28
a

20
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30
is)

22
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Sl 77
=|

15
“T- 20
eT 72
*I70
a

ve €% G le OF 6L St LL OL
5] | al = 3] al eal a sl 2

Fig. 23.—Diagram showing times taken by a kitten to escape from the
cage in twenty-four successive experiments.

successful movement, and gradually associates it with the sense-
impression of the interior of the box until the connection is perfect,

SOME RESULTS OF EXPERIMENT 151

so that it performs the act as soon as confronted with the sense-
impression. . . . Previous experience makes a difference in the quick-
ness with which the cat forms the associations. After getting out of
six or eight boxes by different sorts of acts, the cat’s general tendency
to claw at loose objects within the box is strengthened and its tendency
to squeeze through holes and bite bars is weakened; accordingly it
will learn associations along the general line of the old more quickly.
Associations between licking or scratching and escape are similarly
established, and there was a noticeable tendency to diminish the act
until it becomes a mere vestige of a lick or scratch. After the cat
gets so that it performs the act soon after being put in, it begins to do
it less and less vigorously. The licking degenerates into a mere quick
turn of the head with one or two motions up and down with tongue
extended. Instead of a hearty scratch, the cat waves its paw up and
down rapidly for an instant.”

Such experiments carried out on a different method give
results in line with my own. The conditions are, however,
somewhat unnatural, which I regard as in some respects a
disadvantage. But we need experiments on different methods
—the more the better,—and if the results they furnish are in
accord, their correctness will be rendered the more probable.
It is to be hoped that Dr. Thorndike will devise further ex-
periments in which (1) the conditions shall be somewhat less
strained and straitened, while the subjects are in a more normal
state of equanimity (cannot “ utter hunger ” be avoided ?), and
(2) there shall be more opportunity for the exercise of rational
judgment, supposing the faculty to exist. To establish the
absence of foresight in the procedure of the cats, it is surely
necessary so to arrange matters that the connections are clearly
open—nay, even obvious—to the eye of reason. It appears
that this consideration has not weighed sufficiently with
Dr. Thorndike.

A series of experiments were made to ascertain whether
instruction (in the form of putting the animal through the
procedure requisite for a given act) was in any degree helpful.
The conclusion is that such instruction has no influence.
Those who have had experience in teaching animals to per-
form tricks will probably agree here—though some trainers

152 INTELLIGENT BEHAVIOUR

give expression toa different opinion. It is, however, essential
to distinguish carefully between showing an animal how a trick
is done, and furnishing useful accessory stimuli (such as the
occasional taps of the trainer’s whip when he wants a per-
forming horse to kneel), or affording suitable conditions the
results of which temporarily enter into the association com-
plex. If the latter be eliminated the practice of trainers,
I believe, bears out the general result of the experiments.
Dr. Thorndike never succeeded in getting an animal to
change its way of doing a thing for his. Nor was I, after
repeated trials, able to modify the way in which my dog
lifted the latch of the gate. He did it with the back of his
head. I could not get him to do it (more gracefully) with
his muzzle.

It may be said that the remarkable feats of performing
animals imply the existence of faculties of a higher order than
Dr. Thorndike and I are prepared to admit on the basis of our
experiments. Mr. P.G. Hamerton many years ago described *
how, in his own house, a cleverly trained dog would fetch in
their right order the letters which spelt the English or German
equivalents of common French words, and do other wonderful
things. But the owner of the dog (M. du Rouil) admitted
that there was a means of rapport between them which he was
not prepared to divulge. It is just because the trainer has to
lead up to and utilize chance experiences that such prolonged
patience and care are required. The animal is but the instru-
ment on which his clever trainer plays; an instrument of
wonderful intelligence, but lacking in the higher rational
faculty. The organized scheme is the master’s, not that of
his willing slave. A rational being might not do more
wonderful things; but he would learn them more rapidly
and by a less wearisome method. As it is, the clever per-
forming dog originates little or nothing, and repeats again and
again the same stereotyped behaviour, which—if one witnesses
the performance often—touches one with a profound sense of
its lack of rational spontaneity.

* The Portfolio (1873), p. 27, “ Canine Guests.”

SOME RESULTS OF EXPERIMENT 153

As at present advised, therefore, I see no reason for with-
drawing from the position provisionally taken up. The
utilization of chance experience, without the framing and
application of an organized scheme of knowledge, appears to
be the predominant method of animal intelligence.

On this view, then, we may see in instinctive behaviour,
and the multifarious automatic acts of animals, a means of
providing experience of the right kind and on profitable lines.
We may see in the play-instincts of the young a training
ground for the more serious business of animal life—a theme
developed by Professor Groos. We may see in the imitative
tendency—the innate proclivity to follow a lead blindly and
abt first unintelligently—a further means of providing those
useful items of experience which intelligence finds so service-
able. And we may see in the intelligence which can profit
by chance occurrences that arise in these several ways all that
suffices for the simple needs of animal existence.

With some differences of opinion Dr. Thorndike and I have
much in common in the conclusions to which we have been
independently led as to the method and limits of animal
intelligence. We seem to be in essential agreement in the
belief that the method of animal intelligence is to profit by
chance experience without rational foresight, and that unless
such experience be individually acquired, the data essential for
intelligent progress are absent. While in our attempts to
realize the general nature of animal consciousness there is a
close similarity of treatment. In my “Introduction to Com-
parative Psychology” a good deal of space is devoted to an
analysis of the psychology of skill “in order that we may infer
what takes place in the minds of animals ;” and I said :—
“When I am playing a hard game of tennis, or when I am
sailing a yacht close to the wind in a choppy sea, self does not
at all tend to become focal. Hence,, though I am a self-
conscious being I am not always self-conscious. And pre-
sumably when I am least self-conscious, I am nearest the
condition of the animal at the stage of mere sense experience.
I am exhilarated with the sense of pleasurable existence, my

154 INTELLIGENT BEHAVIOUR

whole being tingles with sentient life. I sense, or am aware
of, my own life and consciousness, in an unusually subtle
manner. Experience is vivid and continuous. Such I take it
to be the condition of the conscious but not yet self-conscious
animal.”

I can therefore cordially endorse Dr. Thorndike’s conclu-
sions as expressed in the following passages :—

“One who has watched the life of a cat or dog for a month
or more under test conditions, gets, or fancies he gets, a fairly
definite idea of what the intellectual [intelligent] life of a cat
or dog feels like. It is most like what we feel when con-
sciousness contains little thought about anything, when we feel
the sense-impressions in their first intention, so to speak, when
we feel our own body, and the impulses we give to it. Some-
times one gets this animal consciousness while in swimming,
for example. One feels the water, the sky, the birds above,
but with no thoughts about them or memories of how they
looked at other times, or xsthetic judgments about their
beauty ; one feels no ideas about what movements he will
make, but feels himself make them, feels his body throughout.
Self-consciousness dies away. Social consciousness dies away.
The meanings, and values, and connections of things die away.
One feels sense-impressions, has impulses, feels the movements
he makes ; that is all.”

And after an illustration from such a game as tennis, Dr.
Thorndike adds : “ Finally the elements of the associations are
not isolated. No tennis-player’s stream of thought is filled
with free-floating representations of any of the tens of
thousands of sense-impressions or movements he has seen and
made on the tennis-court. Yet there is consciousness enough
at the time, keen consciousness of the sense-impressions,
impulses, feelings of one’s bodily acts. So with the animals.
There is consciousness.enough, but of this kind.”

It may be said that between the method of intelligence and
that of fully developed rational procedure there is a wide gap
which must have been bridged in the course of mental evolu-
tion. Unquestionably. And in contending that the methods

EVOLUTION OF INTELLIGENT BEHAVIOUR 155

of the animal are predominantly intelligent, I am far from
wishing to assert dogmatically that in no animals are there
even the beginnings of a rational scheme. Indications thereof
do not indeed at present appear to have been clearly disclosed
by experiment. But the experimental development of the
subject is still in its infancy. We shall probably have to await
the further results which must be the outcome of patient and
well-directed child-study. The human child does pass in the
course of his individual development from intelligent to
rational procedure. Here there is a bridge which is crossed
by every child. When we know more about the stadia of this
development we shall be in a position to apply the results
obtained in child-study in the analogous field of animal-study.
Till then we must possess our souls in patience, and base our
provisional conclusions on the results of systematic investiga-
tion, rather than on those of casual observation and anecdote.

IV.—Tue Evouvution or INTELLIGENT BEHAVIOUR

It is difficult to say where, in the hierarchy of animal
progress, the beginnings of intelligence can first be traced. In
the articulated animals, such as the insects, spiders, and
crustacea, there is abundant evidence of intelligence of a
relatively high grade. Many molluscs unquestionably profit
by experience. The way in which limpets return to the scars
on the rock which form their homes seems to show that they
have acquired a practically adequate experience of their near
surroundings. Romanes cites * some of the earlier observa-
tions which were extended by Professor Ainsworth Davis.t I
looked into the matter myself some years ago, at Mewps Bay
near Lulworth in Dorsetshire. The method adopted f was to
remove the limpets from the rock, and affix them at various
distances from their scars. This can be done without difficulty
or injury to the mollusc if one catches them as they are moving.

* « Animal Intelligence,” pp. 28, 29. t Nature, vol. xxxi., p. 200.
t Ibid., vol. li., p. 127.

156 INTELLIGENT BEHAVIOUR

But one must make sure that they are just leaving or returning
to their proper homes, and are not taken in the midst of a
more extended peregrination, as in that case their special scars
cannot be noted. Failure to be careful in this matter vitiated
my earlier observations, which are therefore excluded in the
following table :—

Number returned.
Number Distance
removed. in inches.
In 2 tides. In 4 tides. Later.

25 6 21 _— _—
21 12 13 5 al
21 18 10 6 2
36 24 1 1 3

From the nature of the rock surfaces the removal of a
limpet to a distance of two feet almost invariably involved
placing them on the further side of an angle. And though
some returned over such an angle, the majority did not.

In most cases the individuals which failed to return to
their respective scars took up new positions; and in several
instances, when they were subsequently removed to a distance
of a few inches from this new position, they returned to it.
Their return to the scar was watched in many cases, and the
course was fairly, but not quite direct. One limpet covered a
distance of ten inches, over a somewhat curved course, in a
little under twenty minutes. In another case the limpet on its
return journey had to pass between two others, which necessi-
tated the lifting of the shell to some height so as to clear one
of them. On reaching the scar they twist and turn about so
as to fit down in the normal position which is constant. When
they come up the wrong way round they rotate pretty rapidly
through the 180 degrees to get into position. One was
observed to make a short excursion from and return to its scar
under stillish water. But as a rule they seem to remain fixed
when they are submerged, moving for the most part when the
tide has just receded.

EVOLUTION OF INTELLIGENT BEHAVIOUR 157

The greatest distance I have watched a limpet reach from
its home was twenty-two inches. But I have found them at a
distance of three feet from their scars—that is to say, from
those to which they fitted perfectly. This was on a large flat
surface.

When they move, the tentacles are projected out beyond
the shell, and keep on touching and slightly adhering to the
rock. On reaching the scar they carefully feel round it with
the tentacles. By excision of these feelers Professor Davis was
led to conclude that it is not through their instrumentality that
the limpet finds its way back to its particular scar. But
I am inclined to question these results. At any rate, further
observations and experiments are needed to settle the point.

Snails will also return to special dark hollows or crannies
in the wall after their foraging excursions. Such behaviour
in molluscs affords evidence of something more than instinct.
In popular speech, we should say that there is memory of the
locality. And in any case it is difficult to interpret the facts
without the assumption that the animals are conscious, and
that re-presentative states are evoked through the mediation of
presentative sense-impressions. And such re-presentative states
are the foundation-stones of experience, which forms the basis
on which intelligent behaviour is grounded.

The most highly developed molluscs are the cephalopods.
They have long sensitive mobile arms with which they feel for
and capture their prey. ‘‘ Now Schneider observed,” writes
Dr. Stout,* “a very young octopus seize a hermit-crab. The
hermit-crab covers the shell in which it takes up its abode
with stinging zoophytes. Stung by these, the octopus imme-
diately recoiled and let its prey escape. Subsequently it was
observed to avoid hermit-crabs. Older animals of the same
species managed cleverly to pull the crab out of its house
without being stung.” Such cases afford evidence of profiting
by experience through the exercise of intelligence.

Darwin’s careful observations on the manner in which
earthworms drag leaves into their burrows seem to show that

* “Manual of Psychology,” p. 257.

158 INTELLIGENT BEHAVIOUR

these annelids act intelligently, and deal with leaves of different
shapes in different ways. The leaves of Pine trees, consisting
of two needles arising from a common base, were almost
invariably drawn down by seizing this basal point of junction ;
while the leaves of the Lime were, in 79 per cent. of the
cases examined, drawn down by the apex; in only 4 per
cent. by the base; and in the remaining 17 per cent. by
seizing some intermediate portion. On the other hand, the
leaves of the Rhododendron, in which the basal part of
the blade is often narrower than the apical part, were in
66 per cent. of the observations drawn down by the narrower
base. Triangles of paper were in the majority of cases
seized by the apex. Commenting upon his observations,
carried out with great care under experimental conditions,
Darwin says,* “ As worms are not guided by special instincts
in each particular case, though possessing a general instinct to
plug up their burrows, and as chance is excluded, the next
most probable conclusion seems to be that they try in different
ways to draw in objects, and at last succeed in some one
way ;” that is to say, they profit by experience based on the
method of trial and failure. But Darwin adds that the
evidence he obtained shows “that worms do not habitually
try to draw objects into their burrows in many different ways.”
And he seems to attribute to them an almost rational power of
dealing with the circumstances in the light of general con-
ceptions. “If worms,” he says, “are able to judge, either
before drawing or after having drawn an object close to the
mouths of their burrows, how best to drag it in, they must
acquire some notion of its general shape. This they probably
acquire by touching it in many places with the anterior
extremity of their bodies, which acts as a tactual organ. It
may be well to remember how perfect the sense of touch
becomes in a man when born blind and deaf, as are worms.
If worms have the power of acquiring some notion, however
rude, of the shape of an object and of their burrows, as seems
to be the case, they deserve to be called intelligent ; for they

* « Vegetable Mould and Earthworms,” p. 95,

EVOLUTION OF INTELLIGENT BEHAVIOUR 159

then act in nearly the same manner as would a man under
similar circumstances.”

Such power of perceiving the relation of the shape of
a leaf or other object to the form of the burrow is presumably
beyond the reach of an earthworm. It may be regarded as
more probable that the earthworm inherits an instinctive
tendency to draw down objects in special ways, and that this
is subject to some modification under the play of experi-
ence, without the formation of anything so psychologically
complex as a general notion, however rude. In any case the
behaviour of earthworms in closing their burrows seems to
afford indications of something more than instinct—of that
profiting by the results of experience which characterizes
intelligent procedure.

Professor Whitman * has made some interesting observa-
tions on the leech Clepsine. “ Place the animal,” he says, “ in
a shallow, flat-bottomed dish, and leave it for a few hours or
a day, in order to give it time to get accustomed to the place,
and come to rest on the bottom. Then, taking the utmost
care not to jar the dish or breathe upon the surface of the
water, look at the Clepsine through a low magnifying lens, and
see what happens when the surface of the water is touched
with the point of a needle held vertically above the animal’s
back. If the experiment is properly carried ont, it will be
seen that the respiratory undulations Gf such movements
happen to be going on) suddenly cease, and that the animal
slightly expands its body and hugs the glass. Wait a few
moments until the animal, recovering its normal composure,
again resumes its respiratory movements. Then let the needle
descend through the water until the point rests on the bottom
of the dish at a little distance from the edge of the body.
Again the movements will cease, and the animal will hug the
glass with its body somewhat expanded. Now push the needle
slowly along towards the leech, and notice as the needle comes
almost in contact with the thin margin of the body, that the
part nearest the needle begins to retreat slowly before it.

* Wood’s Holl Biological Lectures (1898), p. 287.

160 INTELLIGENT BEHAVIOUR

This behaviour shows a surprising keenness of tactile sensi-
bility, the least touch of the water with a needle-point being
felt at once. . . . If its back were rubbed with a brush or the
handle of a dissecting needle, in order to test its sensitiveness
to touch, the appearance would probably be that of insensi-
bility and indifference to the treatment. Closer examination,
however, would show that the flesh of the animal was more
rigid than usual, and that the surface was covered with
numerous stiff, conical elevations, the dermal papillee or warts,
which are so low and blunt in the normal state of rest as to be
scarcely visible. It would be seen that the animal, although
motionless, was in a state of active resistance to attack... .
Clepsine has another and entirely different method of keeping
quiet. The animal rolls itself up (head first and ventral side
innermost) into a hard ball, outwardly passive, free to roll or
fall whithersoever gravity or currents of water may direct
it... . If by chance the animal has eggs, it will not desert
them to escape in this way. . . . This species, then, has two
quite distinct and peculiar ways of keeping quiet, and thus
avoiding its enemies. If the animal has no eggs, or if it has
young, it may adopt either mode of escape, while if it has eggs
it has no choice but to remain quiet over them. . . . The act
of rolling up into a passive ball may be performed (@) under
compulsion, as when it is her last resort in self-defence ;
(0) under a milder provocation, aS one of three courses of
behaviour, as when the resting-place is turned up to light, and
the choice is offered between remaining quiet in place, creeping
away at leisure, or rolling into a ball and dropping to the
bottom ; (¢) or finally, wnder no special external stimulus, but
rather from internal motive, the normal demand for rest and
shady seclusion, presumably very strong in Clepsine after
gorging itself with the blood of its turtle host.”

Professor Whitman rightly regards the act of rolling into
a ball as instinctive, and due to natural selection. But he does
not undertake to discuss the question as to how much intelli-
gence, if any, Clepsine may have. Nor, indeed, is it an easy
matter to determine. The differential reaction according as

EVOLUTION OF INTELLIGENT BEHAVIOUR 161

the animal has eggs or not suggests intelligence; but it may
be instinct varying according to the conditions of stimulation
external and internal. The different behaviour which may be
seen in different cases when a stone is turned to the light
again suggests intelligence, but again may be determined
directly by the conditions of stimulation. Prompted by Dr.
Whitman’s observations, I endeavoured to determine whether a
leech would grow accustomed to frequent gentle stimulation
with a camel’s-hair brush, and cease to react under circum-
stances which were followed by no ill effects. But though I
incline to think that this is the case, the observations were not
such as to be satisfying and convincing. If intelligence be
present we seem to find it in an early and rudimentary state.

Observation and experiment seem to afford little indication
of the conditions under which intelligence first makes its
appearance in the animal kingdom. And if we turn to
general considerations, which at the best afford uncertain
guidance, little light is thrown on the subject. If we accept
the view already indicated,* that the nerve-centres which are
concerned in the conscious control begotten of experience are
independent of those primarily concerned in normal reflex
action, we may perhaps believe that the simplest nervous
system, worthy of the name, contains both these elements, and
that in the course of the evolution of nervous systems in
higher and higher grades, there go on part passu the further
differentiation of these elements, and the progressive integra-
tion of reflex and control centres into a closely connected and
effective whole. Not that any expression of the facts, if such
they be, in terms of an evolution formula, adds anything to
our knowledge of the organic modus operandi. We know
but little of the intimate nerve physiology of even the highest
invertebrates. We see ample evidence of the control of
behaviour in the light of individual experience. Of any
detailed knowledge concerning the manner in which this
control is effected we do not seem to possess more than the
rude initial phases.

* Supra, p. 44.

162 INTELLIGENT BEHAVIOUR

When we compare, however, the several grades of intelli-
gence which observation discloses, and when we watch the
conscious development of the more intelligent animals, we seem
to find evidence of the growth of a system of experience, at
first in very close touch with inherited modes of procedure,
but gradually acquiring more of independence and freedom.
Increase of the range and complexity of behaviour brings with
it, not only increase in the range and complexity of experience,
but also—what is, perhaps, even more essential to effective
progress—greater unity and closer connection into a well-knit
whole. And with this greater unity and closer connection
there goes what one may term a condensation of experience by
an elimination of detail and the survival of essential features
repeatedly emphasized. This is analogous in the development
of intelligence to the generalization and abstraction which play
so important a part in the development of reason. It affords,
in fact, the data which reflection utilizes in the purposive and
intentional condensation and concentration of knowledge at a
higher stage of mental development.

The omission of detail and the survival of the salient
features is well known to us in the familiar facts of memory.
We have seen thousands of sheep and oxen, no two of which
are probably alike in all their external details as presented to
vision. But we remember what a sheep or an ox looks like,
and many of us can form a visualized image of either of these
animals, This, however, is not the re-presentative image of any
particular sheep or ox. It is what psychologists term a generic
image. It is like a composite photograph made by superimpos-
ing on the same plate a number of individual images so that the
salient features which all possess in common stand out clearly
by their coincidence on the plate, while the distinctive details
are but dimly presented. Thus does memory preserve the
essentials common to many impressions while the distinguishing
details are lost and fade, eliminated by forgetfulness. And
thus in the experience which intelligence practically utilizes
are the net results of a thousand particular impressions con-
densed in one effective image.

EVOLUTION OF INTELLIGENT BEHAVIOUR 163

Condensation of experience is also effected by the elimina-
tion, under the guidance of consciousness, of those modes of
behaviour which are not efficacious—a process to which Pro-
fessor Mark Baldwin applies the phrase Functional Selection.
There is a tendency at first to the overproduction of relatively
useless actions. The multifarious random movements of the
human infant, though their inexactness renders the child
terribly helpless, afford a wide store of plastic material which
intelligence can guide to its appropriate use. And the pro-
longed period of pupilage in the child is correlated with an
unsurpassed range of combination and recombination of the
abundant plastic material. The hereditary legacy, though it
contains fewer drafts for definite and specific purposes than are
placed to the credit of an animal rich in instinctive endowment,
affords a far larger general fund on which intelligence may
draw for the varied purposes of the freer financial existence
of a rational being.

The relatively helpless young of many of the higher
mammalia exhibit also much overproduction of seemingly
aimless movements. But from these intelligence selects those
which are of value for the purposes of life—those which
experience proves to be effective. These—the relatively few—
afford the motor impressions which by repetition stand out in
experience, while the rest lapse from memory and are elimi-
nated from experience as they are eliminated from practical
performance. This is a great gain. Motor experience is
rendered generic; the composite image that is retained is the
net result of effective behaviour; and all that is valuable in
the acquisitions of early life is condensed within manageable
limits.

This process of rendering generic the particular items of a
widening experience has a marked effect in the development of
the conscious situations in the light of which behaviour is
intelligently guided. It is not the master holding this whip
or that ball which suggests to the dog a hiding or a scamper ;
it is a generic situation with interchangeable details. It is not
this, that, or the other previously unseen cat that at once

164 INTELLIGENT BEHAVIOUR.

determines the situation for the fox terrier; the particular
animal has never entered into his past experience: it is the
fulfilment of the essential conditions of the generic image that
is operative in behaviour. The experience of animals must
inevitably become in large degree generic by the elimination
of the unessential and survival in re-presentative consciousness
of the salient elements in many slightly diverse situations.

Stated in terms of this conception, the familiar phenomena
of mimicry are due to the fact that the mimicking form accords
sufficiently well with the generic image to be taken as a
representative thereof. As is well known, the model has been
proved in many cases to be unpalatable or hurtful, while the
mimic is in itself neither the one nor the other. The drone-
fly, Hristalis, mimics the drone. And it has been urged that
this cannot be a true case of mimicry, since the drone is harm-
less, though the female and “neuter” bees are possessed of stings.
But I have satisfied myself by experiments with young birds, that
(1) after experience with bees drones are avoided, and (2) that
after similar experience drone-flies are also left untouched.
Hence it seems that all three fall within the same generic image,
the points of resemblance outweighing the differences in detail
—as they do, indeed, with many men and women.

Such examples of mimicry belong to what is known as the
“Batesian type ”"—so called after HL. W. Bates, who, in 1861,
discussed its occurrence among Amazonian insects in the light
of the theory of natural selection. There are, however, certain
groups of insects which, although themselves ‘“ protected,”
possess common warning colours, causing them to resemble
each other. These are sometimes classed under the head of
“‘Mullerian mimicry ”’—so called after Fritz Miller, who, in
1879, first offered an explanation of the facts based on the
theory of natural selection. He suggested that such mutual
resemblance is advantageous to both protected forms, since it
lessens the number of those which are killed by young birds
and other animals while they are learning by experience what
to eat and what to leave. For, as the result of careful
observation, Mr. Frank Finn concludes ‘that each bird has

EVOLUTION OF INTELLIGENT BEHAVIOUR 165

to separately acquire its experience, and well remembers what
it has learnt,’”—a conclusion with which, as already stated, my
own observations are entirely in accord. There is therefore a
certain amount of destruction of even well-protected forms
by young and inexperienced birds. If, then, two such forms
resemble each other, the acquisition of experience is thereby
facilitated and the amount of destruction reduced, on the
assumption that the two fall within the same generic image.
Upholders of natural selection are not, indeed, at one in accept-
ing this explanation, and further observation is unquestionably
needed. It is not improbable, however, that common protective
coloration, such as the banding of yellow and black, seen in such
different forms as the caterpillar of the cinnabar moth and the
imago of the wasp, is of mutual utility. The following experi-
ment was made with young chicks. Strips of orange and black
paper were pasted beneath glass slips, and on them meal
moistened with quinine was placed. On other plain slips
meal moistened with water was provided. The young birds
soon learnt to avoid the bitter meal, and then would not touch
plain meal if it were offered on the banded slip. And these
birds, save in two instances, refused to touch cinnabar cater-
pillars, which were new to their experience. They did not,
like other birds, have to learn by particular trials that these
caterpillars are unpleasant. Their experience had already been
gained through the banded glass slips; orsoitseemed. I have
also found that young birds who had learnt to avoid cinnabar
caterpillars left wasps untouched. Such observations must be
repeated and extended. But they seem to show that one aspect
of the Miillerian theory is not without some facts in support of
it; and, so far as they go, they afford evidence that black and
orange banding, irrespective of particular form, may constitute
a guiding generic feature in the conscious situation.

It may be said that the generic condensation of experience
here indicated implies the formation of general and abstract
ideas, and that we cannot in face of the evidence accept
Locke’s dictum that abstraction is ‘an excellency which the
faculties of brutes do by no means attain to.” Romanes

166 INTELLIGENT BEHAVIOUR

contended * that “all the higher animals have general ideas of
‘eood-for-eating ’ and ‘ not-good-for-eating ’ quite apart from
any particular objects of which either of these qualities happens
to be characteristic,” and he quoted with approval Leroy’s
statement, that a fox “ will see snares when there are none ;
his imagination, distorted by fear, will produce deceptive
shapes, to which he will attach an abstract notion of danger.”
According to such views animals form concepts ; and concepts
belong to the sphere of rational thought. It is not my inten-
tion to enter at length into the refinements of psychological
distinction. Many psychologists, however, seek to distinguish
between, on the one hand, the predominance by natural
emphasis, of certain qualities, such as that of being suitable for
food, and, on the other hand, the intentional isolation of these
qualities for the purposes of thought and rational explanation.
Abstraction they regard as a deliberate process applied with
rational intent to the material afforded by experience and
reflection. Generalization, too, they regard as deliberate, and
carried out with like intent. The result is not merely a
composite or generic product, but something more subtle and
less dependent on sense. “All trees hitherto seen by me,”
said Noire, “leave in my imagination a mixed image, a kind
of ideal presentation of a tree. Quite different is my concept,
which is never an image.” The concept “tree ” is a deliberate
synthesis of abstract qualities intentionally isolated, and
recombined in accordance with the general relationships which
subsist between them.

If we accept this distinction, if we regard abstraction and
generalization as intentional mental processes carried out with
the rational intent of discovering the relationships of pheno-
mena with the object of explaining them and recombining
their essential features in an ideal scheme of thought, we shall
probably admit, with John Locke, that these are excellencies
which the faculties of brutes do by no means attain to. But
we shall none the less see that the predominance of certain

* « Mental Evolution in Man,” p. 27.
+ “Intelligence of Animals,” p. 121

EVOLUTION OF INTELLIGENT BEHAVIOUR 167

salient features in experience by reiterated emphasis in
association with natural needs, and the development of generic
in place of merely particular re-presentations will afford the
appropriate material for abstraction on the one hand, and
generalization on the other. Intelligence supplies the em-
bryonic mental structures from which, under the quickening
influence of a rational purpose, abstract and general ideas may
be evolved.

The essential features of the evolution of intelligence seem,
then, to be, first, the development of controlling nerve-centres,
by which the responsive action of reflex automatic or instinc-
tive centres may be checked, augmented, or modified ; secondly,
the increased differentiation and integration of these control
centres with extension of the range and complexity of experi-
ence in close touch with practical needs; thirdly, the con-
densation and concentration of experience by the formation of
generic products through the reiterated emphasis begotten of
recurrent situations having certain salient features in common,
though differing in details; and fourthly, an increased
plasticity of behaviour, especially in early life, enabling an
animal to deal effectually with an environment more com-
plicated than that to which the more stereotyped instinctive
behaviour is fitted by inheritance to respond. And this
evolution of intelligent behaviour is working its way up to,
though as such it cannot reach, the succeeding phase of mental
evolution in which the data, supplied by intelligence, are
treated with a new purpose for higher ends in the rational
thought which seeks to explain the phenomena, and frame an
ideal scheme of their relations and interconnections.

Two further points may be noticed. First, that it is during
the early and plastic days or months of life that intelligence is
setting its seal on animal behaviour, and stamping it with its
distinctive character. Adult life is very much what youth has
made it; and old age is stereotyped through habit. In times
of progress, the character of the race is determined by plastic
possibilities of the young. Among them it is that the incidence
of elimination makes itself felt, resulting in the survival of those

168 INTELLIGENT BEHAVIOUR

whose intelligence can mould behaviour in accordance with
the new circumstances of a wider life.

Secondly, this selection of the intelligent involves the
survival of those in whose higher brain-centres there is room
for a greater range and variety of interconnection by means of
associating fibres. It involves a selective survival of the
larger and more finely organized brains. It is probable, as
Professor Ray Lankester has recently indicated, that the
ridiculously small-brained mammals and reptiles of the past
were creatures of instinct with little capacity for intelligent
control. Their lives were simple, and their enemies and com-
petitors no better provided with higher brain-centres than
themselves. Stereotyped instinctive behaviour sufficed to
enable them to hold their own, and meet the requirements of
a life of dull and unprogressive monotony. Strength without
cunning made these big-framed animals for a while masters of
the situation. But there are no existing animals, whose
skeletons indicate so high a position in the zoological series,
which exhibit a cerebral development so poor. And we may
fairly conclude that the fact that these huge creatures have
left no lineal descendants may be taken as evidence of the
importance and value, in evolution, of that cerebral tissue
which is the organic basis of intelligence. The higher brain
contains the potentiality of that experience without which
the evolution of intelligent behaviour in any race of vertebrate
animals is impossible.

V.—TuHE INFLUENCE OF INTELLIGENCE ON INSTINCT

We have seen thai the relation of instinct to intelligence is
essentially that of congenital to acquired behaviour. We have
seen, too, that in the Lamarckian interpretation what is
acquired in the course of life may be transmitted through
inheritance, and thus the intelligent behaviour of one genera-
tion may become instinctive and congenital in the next. But
serious biological difficulties stand in the way of the acceptance

INFLUENCE OF INTELLIGENCE ON INSTINCT 169

of this interpretation ; there is, moreover, little or no evidence
of the assumed transmission to offspring of any acquired
modifications of structure or behaviour. We have, therefore,
been led to infer that instinctive behaviour has been evolved
through the selection of adaptive variations of germinal origin,
the influence of intelligence being restricted :to the fosterage
of co-incident variations, that is to say, of those congenital
variations which coincide in direction with the acquired
modifications of behaviour due to intelligence. It is clear
that on this interpretation the influence of intelligence on
instinct is more indirect and less simple than that implied by
the Lamarckian hypothesis. Intelligence and instinct are in
large degree independent, though there is continual interaction
between them. We have now to consider the nature of this
interaction, and to this end we must indicate the relation of
acquired modifications to the hereditary groundwork of the
animal constitution.

The basal fact is, that the bodily tissues are subject to a
certain amount of structural change during the course of
individual life in accordance with the amount of functional
strain put upon them. The labourer’s thickened skin, the
enlarged and strengthened muscles of the athlete, the jugeler’s
acquired suppleness are familiar cases. Less familiar instances
are afforded under abnormal conditions. Should one kidney
from any cause be slowly destroyed, the other will slowly
enlarge to carry on the increased work of elimination of waste
products; when the larger shin bone of a dog has been
removed after injury, the smaller bone becomes thickened to
bear the added strain ; new joint surfaces are sometimes formed
where bones have been broken and the natural joints injured.

One may say that the normal development of any structure
depends upon a due amount of use. But, since in the course
of strenuous life any organ is from time to time subject to an
abnormal amount of strain, it must be fitted to respond to a
super-normal call on its strength and functional activity.
Were the heart and the lungs, for example, unable to meet the
greatly added drain on their energies, due to unwonted and

170 INTELLIGENT BEHAVIOUR

severe exertion, collapse, perhaps death, would ensue if such
exertion were imperatively demanded under special circum-
stances. And it is clear that many wild animals must be not
infrequently placed in such circumstances as will subject their
muscular structures and the functional activity of their organs
of circulation and respiration to a strain nearly up to their
extreme limits of endurance. The carnivorous hunter would
often fail to secure his prey if his organization were unequal to
a hard and prolonged chase; the hunted prey would not
survive to procreate his kind if he fell a victim to the first
pursuer through inability to stand the exertion necessary to
enable him to make good his escape. It is thus, we may
believe, through natural selection that a sufficiently high
standard of streneth and functional endurance is maintained.
The failures in these respects are steadily eliminated. It is
difficult to realize the great strain put upon a bird’s organization
by the migration flight. Some ten times as many birds leave
our shores in the autumn as return to them in the following
spring. What proportion of these is weeded out in the act of
migration we do not know; but we may be sure that only
those fitted to stand a severe test of physical endurance return
to rear broods which shall inherit in large degree similar vigour
of constitution.

Two factors, then, determine the limits of efficiency in the
bodily organs—heredity and use. And these two co-operate in
such a way that we may say, either that due use is the essential
condition of the effective development of the hereditary powers,
or that heredity serves to condition their effective development
through use. But though closely related, so that each may be
regarded as conditional on the other, they are, if we accept the
view that acquired characters are not transmitted as such, so
far independent in that use adds nothing to, disuse subtracts
nothing from, the hereditary store. It is, indeed, difficult
to conceive how, on any view, the absence of the conditioning
factor of normal use can be the efficient cause of a positive
diminution of the balance at the bank of heredity. And
Lamarckian thinkers have not succeeded in placing their

INFLUENCE OF INTELLIGENCE ON INSTINCT 171

conception of the matter in the clear light of a working
hypothesis.

The amount of what we may term “ modifiability ” by use
differs a good deal in the several organs and tissues. The
teeth of carnivora and the antlers of deer may be cited as
structures in which the conditioning effects of use form a
relatively unimportant factor. On the other hand, the nervous
system, with which we are here primarily concerned, is of all
animal structures that in which what is acquired may attain
the greatest importance in the successful conduct of life; the
nature and the range of behaviour affording an index of the
amount of modifiability in this respect.

We have already seen that instinctive behaviour is primarily
a matter of the first occasion on which any given action is
performed, and that many instinctive acts are subject to sub-
sequent modification in the light of the experience gained
during the early performances. The range of such modification
varies both in different animals and also with respect to
different modes of behaviour in the same animal. The more
fixed and deeply rooted an instinct the less readily does
intelligence obtain a hold on it, so as to direct the behaviour
into new channels of better accommodation to the circum-
stances. M. Fabre describes how a Sphex, one of the solitary
wasps, instinctively draws its prey, a grasshopper, into the
burrow by its antennee. When these were cut off the wasp
pulled the grasshopper in by the jaw appendages ; but when
these were removed she seemed incapable of further accom-
modation to the unusual circumstances. It would seem an
easy and obvious application of intelligence to seize the prey by
one of the forelegs. But this was not done; and the grass-
hopper was then left. Intelligence did not seem equal to
meeting the altered conditions presented by the maimed grass-
hopper. Still, there was some modification of the normal
instinctive behaviour ; and, as Dr. Peckham has shown, there
may be more than Fabre noted. Let us assume the exist-
ence of an animal whose every act is instinctive, whose
whole behaviour is marked out in strictly hereditary lines, no

172 INTELLIGENT BEHAVIOUR

new departures being acquired in the course of individual life.
This extreme case would afford an example of what we may
term completely stereotyped behaviour. On the other hand, let
us assume the existence of an animal with no hereditary
definiteness of reaction, whose every act is intelligent, whose
whole behaviour is the result of individual acquisition. This
antithetical extreme case would afford an example of what we
may term completely plastic behaviour. It is questionable,
however, whether either of these extreme types occur in nature.
What we find in our study of animal behaviour is some inter-
mediate condition in which both factors co-operate, with a
predominance either of stereotyped instinctive response on the
one hand, or of plastic intelligent acquisition on the other
hand. And in the latter case, as such behaviour approaches its
ideal limits, we have modifiability under the circumstances of
individual life at its maximum.

The evolution of intelligence as such runs parallel with the
evolution of plastic behaviour ; and this plasticity is necessi-
tated by the variety and the complexity of the conditions of
life—a variety and a complexity requiring many subtle modifi-
cations of response to enable the behaviour to reach accommoda-
tion to the changeful exigencies of diverse circumstances. To
meet constant and relatively fixed conditions stereotyped
instinctive responses suffice ; and the elimination under natural
selection of those individuals which fail to respond in fixed
ways by specially adaptive behaviour tends to render definite
the hereditary channels of nervous intercommunication. An
inherited system of no little complexity may thus be evolved ;
of which we have seen examples in our study of instinctive
behaviour. But the essential condition of the successful
working of such a system is unvarying constancy in the
environment to which the stereotyped instinctive behaviour is
adapted. Instinct without intelligence is like a barrel-organ
constructed to play a limited number of tunes with monotonous
precision. The music of its behaviour depends entirely on
what the maker, heredity, has inserted in the works. But
though a barrel-organ may suffice where a hymn-tune, a jig, a

INFLUENCE OF INTELLIGENCE ON INSTINCT 173

hornpipe, and a funeral march exhaust all the possible require-
ments, it is sadly lacking in musical plasticity. To obtain
that, you must place intelligence at the keyboard that the music
may be accommodated to a greater number of varying moods.

It may be urged that such an illustration is, in many
respects, obviously faulty ; and that barrel-organs do not under
any circumstances develop into musicians. No doubt the
illustration is faulty. But it may be questioned whether
instinct under any circumstances develops into intelligence, any
more than a barrel-organ into a musician. As we said at the
beginning of this section, intelligence and instinct are in large
degree independent though there is continual interaction
betweén them. Completely stereotyped behaviour, in its
theoretical perfection, is in exact adaptation to the circum-
stances. Where instincts are only relatively perfect, further
adaptation is secured through congenital variation and the
survival of the individuals in which the behaviour is better
adapted to the comparatively invariable circumstances. This
is one line of evolution. But it no more contains within itself
the potentiality of developing into plastic accommodation to
varying circumstances, than the barrel-organ contains within
itself the potentiality of becoming a musician. The evolution
of intelligence is along independent lines of progress. Stereo-
typed adaptation can never pass up into plastic accommodation.
These two belong to independent lines of evolution ; but they
are bound up in the same nervous system, they jointly determine
the behaviour, they interact not only in the course of individual
life but in the process of evolution, and they are both subject
to the incidence of natural selection, which can determine
whether the one line or the other shall preponderate—whether
instinct or intelligence shall dominate behaviour.

If an answer must be given to the question whether instinct
or intelligence has priority in the course of the evolution of
hehaiour, it may be urged that, on theoretical :grounds, the
claims of instinct are the stronger. No doubt the evolution
of the two lines of development have proceeded to a large
extent side by side ; but whereas intelligence does in a number

174 INTELLIGENT BEHAVIOUR

of cases demonstrably modify the course of instinctive behaviour,
the converse proposition does not hold good in the same sense.
The animal acts at first instinctively, and subsequently in the
light of experience reaches further accommodation ; and though
in later life a dominating instinct may override the guidance
of intelligence, still, even this is probably due to the fact that
the instinct is more deeply seated in the constitution than any
opposing habits of intelligent acquisition.

We can, however, infer what is the influence of intelligence
on instinct without basing our inferences on any assumption
of the initial priority of instinct in the evolutionary sequence.
Taking animals as we find them, they afford numberless ex-
amples of behaviour at first instinctive but subsequently modi-
fied, in greater or less degree, in accordance with the teachings
of experience. Let us, first, assume that the environment is
slowly changing, or has changed, in some definite manner.
Such change would, of course, be relative, and might be due,
either to new conditions brought to bear on the animal, or to
the animal being itself brought, in the expansion of its life,
within their influence. The old instinct is no longer quite
adapted to the changed circumstances. If the change were
sufficient in amount, and occurred somewhat suddenly, variations
of instinct might not occur soon enough to enable the animal
to reach adaptation by the gradual process of natural selection.
If dependent on instinct alone the animal would, under these
circumstances, be eliminated. But if intelligence were able to
modify the behaviour to meet the new conditions this elimina-
tion would be prevented. In successive generations intelligence
would constantly modify behaviour in the same manner and in
a definite direction. Meanwhile congenital variations in
different directions would occur. Those which were in direc-
tions antagonistic to that dictated by intelligence would tend
to thwart accommodation and render it less effectual ; but
those which were coincident in direction would conspire with
accommodation and render it more effectual. The individuals
in which variations of instinct tended to thwart intelligence
would be eliminated ; while those in which coincident variations

INFLUENCE OF INTELLIGENCE ON INSTINCT 175

assisted and aided intelligent modification would survive.
- Thus intelligence would lead the way along lines which con-
genital variations would follow. And in the course of a number
of generations the new instinct would reach the fully adaptive
level, and further modification by intelligence would become
unnecessary unless the environment continued to change yet
more. Individual accommodation of behaviour would in this
way determine the direction of instinctive variation ; and yet
throughout the process there would be, strictly speaking, no
transmission of the intelligently acquired characters of the
behaviour.

But though under constant and uniform changes in the
environment the net result would be only a guided variation
of the original instinct, under more variable and indefinitely
changing circumstances the result would be different. The
higher animals exhibit an intelligent plasticity which enables
them to meet the requirements of the more complex environ-
ment into which their wider life has risen ; for evolution lifts
the animal from narrower into progressively wider spheres of
activity and behaviour, so that its environment becomes
relatively more complex. Here stereotyped behaviour would
be rather a hindrance than an advantage. The winning
animal in life’s struggle would be the one in which behaviour
was most rapidly and most surely modified to meet particular
needs—the one in which the teachings of experience were most
promptly utilized in effective action. The inevitable tendency
of the evolution of intelligence must be disintegration of the
stereotyped modes of behaviour and the dissolution of instinct.
Natural selection, which under a uniform and constant
environment leads to the survival of relatively fixed and
definite modes of response, under an environment presenting
a wide range of possibilities leads to the survival of plastic
accommodation through intelligence. It is not that intelligence
has any direct influence tending to undermine the hereditary
foundations of instinct, for acquired plasticity is not inherited
as such ; it is rather that when the stereotyped and the plastic

are pitted against each other in the struggle for existence in

176 INTELLIGENT BEHAVIOUR

the wider, freer, and more varied life of the higher animals
the plastic survives and the stereotyped succumbs.

Imperfect as is our present knowledge of the manner
in which the nervous connections implied in psychological
associations are established, there can be no question that
they are acquired in the course of individual life; they are
modifications of nervous structure due to a special mode of
use under the conditions of experience. Here, then, in the
case of the nervous system, as in that of the bodily organs
before mentioned, two factors determine the limits of efficiency
—heredity and use. And these two, again, co-operate in such
a way that we may say, either that due use—here represented
by adequate experience—is the essential condition of the
effective development of the hereditary potentialities, or that
heredity serves to condition their effective development
through use and experience. And just as the heart and lungs
must inherit the power of standing abnormal strain if the
animal is to avoid elimination in times of unwonted exertion,
so must the nervous system inherit some reserve power of
dealing effectively with unwonted circumstances by intelligent
accommodation, if the animal is not to fall a victim to such
circumstances. In other words, at times of heightened com-
petition those animals which can draw on a reserve fund of
intelligent accommodation will survive, while the stupid
blunderers will be eliminated. We may term this reserve
fund of intelligent accommodation, this inherited ability to
meet specially difficult circumstances as they arise, inate
capacity. From the nature of the case it must be indefinite,
for it must carry with it the ability to meet unforeseen
combinations of the environing forces by new combinations
of the results of experience. Its distinguishing mark is
plasticity, in contradistinction to the stereotyped fixity of
typical instinct. And accompanying its evolution there is
probably, as we have seen, a dissolution of its antithesis,
instinct. Thus may we account for the fact that man, with
his great store of innate capacity, has so small a number of
stereotyped instincts.

INFLUENCE OF INTELLIGENCE ON INSTINCT 177

But the dissolution of instincts is not complete. Residua
are left in the inherited mental constitution. And these we
term congenital tendencies and propensities. They differ from
the typical instincts in the fact that the definiteness of response
has been lost. They dictate a general trend of action, but
the particular application in behaviour is due to intelligent
accommodation. They are commonly spoken of as instinctive ;
and their mode of origin justifies the use of the adjective in
association with the term “ propensities.” But it must be
remembered that the behaviour to which they lead is not, as
such, wholly instinctive ; it is a joint product of instinct and
intelligence, the general trend being due to the instinctive
propensity, while the mode of application is guided by
intelligence.

There is, however, another way in which analogous pro-
peusities may be ingrained in the mental constitution. It is
a well-known and familiar fact that the frequent repetition
of intelligent accommodation in certain definite lines begets
habits, which so far simulate instincts as to be commonly
described in popular speech as instinctive. Professor Wundt
indeed places them in the category of ‘acquired instincts ”»—
a usage which we regard as unsatisfactory, seeing that it tends
to mask the distinction between the congenital and acquired
factors in behaviour, and seeing that we have the well-defined
term “habits” for acts rendered to a large extent automatic
through repetition. Lamarckian thinkers regard habit as the
mother of instinct, assuming that the acquired automatism of
one generation may be transmitted to become congenital in
the succeeding generation. This conclusion we provisionally
reject regarding the basal assumption as at present unproven.
But though we cannot accept the view that habit is the mother
of instinct, we regard it as not improbable that habit may
be the nurse of congenital propensities. Remembering that
similar habits are acquired by animals of the same species
throughout a series of succeeding generations, and assuming
that congenital variations are constantly occurring in many
directions, it seems probable that some of these variations will

N

178 INTELLIGENT BEHAVIOUR

be coincident in direction with the acquired habits. Thus
would arise a congenital propensity to perform the habitual
acts; and should they be of sufficient importance in the
conduct of life to be subject to the action of natural selection,
those animals in which such propensities were congenital
would survive, whereas those in which no such propensities
existed would be eliminated. It is unnecessary, however, to
elaborate this conception further, since it is in line with that
already discussed in considering the influence of intelligence in
fostering a diversion of instinct under changing circumstances.

Sufficient has now been said to illustrate some of the ways
in which instinct and intelligence interact in the evolution of
behaviour. Such interaction is further exemplified in the
social life of animals, which will be dealt with in the next
chapter.

CHAPTER V
SOCIAL BEHAVIOUR

I.—Imitatron

THE characteristic feature of social behaviour is that it is in
large degree determined by the behaviour of other members of
the social community. In all animals which mate there is a
temporary or more lasting influence on each other of the
individuals which unite to procreate their kind ; and in those
which foster their young there is a social relation of parents
and offspring. Some of these mutual relationships will be
discussed, in their emotional aspects, in the next chapter.
Here we will consider the more general factors which serve to
determine the course of social evolution.

Among these is commonly reckoned imitation. M. Tarde
says, “La socicté c’est limitation.” But this word, like so
many others which are employed alike in popular speech and
in more or less technical discussions, carries a somewhat wide
range of meaning, and is by some writers used in a broader, by
others in a narrower sense. Thus Professor Mark Baldwin *
says, ‘‘ that all organic adaptation in a changing environment
is a phenomenon of biological or organic imitation,” under which
category will fall, therefore, the organic behaviour of the
protozoa and of plants, On the other hand, Professor HK. L.
Thorndike, though he admits in the lower animals “ certain
pseudo-imitative or semi-imitative phenomena,” has been led
by experiments, to be presently noticed, to the conclusion that

* “Mental Development in the Child and the Race— Methods and
Processes,” p. 278.
179

180 SOCIAL BEHAVIOUR

animals as high in the scale of life as cats and dogs cannot
form new associations under the influence of imitation. “It
seems sure,” he says,” “from these experiments, that the
animals were unable to form an association leading to an act
from having seen another animal, or animals, perform the act
in a certain situation.” In face of such apparently diverse
usage it is necessary to show within what limits and with
what qualifications the word may profitably here be used to
indicate a factor in social evolution.

Professor Mark Baldwin’s use of the term “imitation ” can
only be understood in its relation to an hypothesis of organic
and mental evolution, which he develops with no little skill
and brilliancy. f He regards the processes of life as issuing in
a great twofold adaptation, due to expansions and contractions,
—the former representing waxing, the latter waning vitality ;
and he holds that all special adaptations are secured by the new
hold upon beneficial stimulations reached by the expansive out-
reaching movements. ‘‘ Among the variations in organic
forms,” he says, “it is easy to see that some of them might
react In such a way as to keep in contact with the stimulus, to
lay hold of it, and so keep on reacting to it again and again—
just as our rhythmic action in breathing keeps the organism in
vital contact with the oxygen of the air. These organisms will
get all the benefit or damage of the repetition or persistence of
the stimulus, or of their own reactions, again and again ; and
it is self-evident that the beneficial stimulations are the ones
which should be maintained in this way, and that the organisms
which did this would live. The organisms which reacted in
such a way as to retain the damaging stimulations, on the other
hand, by this same process, would aid nature in killing them-
selves. If this be true, only those organisms would survive
which had the variation of retaining useful stimulations in
what I have called, in speaking of imitation elsewhere, a ‘ circular
way’ of reacting... . So, when we come to consider phylogeny

* “Animal Intelligence:” monograph supplement to Psychological
Review, 1898, p. 61.
t Op. cét., pp. 263, 172, 201, 132, and 248 (note).

IMITATION 181

and ontogeny together, we find that if by an organism we
mean a thing of contractility or irritability, whose round of
movements is kept up by some kind of nutritive process sup-
plied by the environment— absorption, chemical action of
atmospheric oxygen, etc.—and whose existence is threatened
by dangers of contact and what not, the first thing to do is to
secure a regular supply to the nutritive processes, and to avoid
these contacts. But the organism can do nothing but move,
as a whole or in some of its parts. So, then, if one of such
creatures is to be fitter than another to survive, it must be the
creature which, by its movements, secures more nutritive pro-
cesses and avoids more dangerous contacts. But movements
toward the source of stimulation keep hold on the stimulation,
and movements away from the contacts break the contacts ;
that is all. Nature selects these organisms; how could she do
otherwise ?”

“Thus a ‘circular’ activity is found in operation ; life-
processes issuing in increased movements, by which in turn the
stimulations to the life-processes are kept in action.” But
when a child imitates, himself reproducing the ‘‘ copy ” set for
imitation, the reaction at which imitative suggestion aims is
one which will reproduce the stimulating impression, and so tend
to perpetuate itself. The stimulus starts a motor process,
which tends to reproduce the stimulus, and, through it, the
motor process again. It isa “circular activity.” ‘hus “we
are able to reconstruct the theory of adaptation in such a way
as to show that this kind of organic selection by movement,
and this kind of imitative selection by consciousness, are the
same thing. Organic imitation and conscious imitation—each
a circular process tending to maintain certain stimulations and
to avoid others—here is one thing;” and to this one thing the
common term “imitation” is applied by Mr. Baldwin.

This extended usage is admitted by the author to be some-
what of an innovation. But if his hypothesis be sound this
need be no bar to its acceptance. Two salient questions must,
however, receive satisfactory answers. First, is all organic
adaptation in a changing environment a circular process—a

182 SOCIAL BEHAVIOUR

phenomenon of organic imitation? Secondly, does all con-
scious imitation tend to reproduce the imitating stimulus ?

Professor Baldwin speaks of organic imitation and conscious
imitation as “each a circular process tending to maintain
certain stimulations and to avoid others.” Now, it may be
granted that the tendency to maintain or repeat certain stimu-
lations may be regarded as a “circular process.” But can the
avoidance or non-repetition of others be so regarded? A large
proportion alike of the hereditary adaptations and the acquired
accommodations of behaviour are directed to this avoidance or
non-repetition of hurtful stimulations. The instinctive shrink-
ing of a chick from an aggressive animal is just as much
adaptive as the repeated cuddling beneath the warm wing of
the mother. The avoidance of nauseous cinnabar caterpillars
is just as much an accommodation to the constitution of the
environment as the reiterated seizing of palatable grubs. Even
low down in the scale of animal life, Dr. Jennings’s observations
on Paramecia seem to show that the retention of favourable
stimulation is not due to its direct influence, but is the indirect
result of a reaction to the relatively unfavourable stimulation
which occurs when the Paramecium passes away from more
satisfactory surroundings. A favourable environment is secured
through the avoidance of the unfavourable. Unless, therefore,
we exclude adaptive avoidance from the category of adaptations,
we cannot regard all organic adaptation in a changing environ-
ment as a phenomenon of organic imitation due to a circular
process tending to the reinstatement of stimulation.

Passing to the second question—Does all conscious imitation
tend to reproduce the initiating stimulus ?—we cannot un-
reservedly give an affirmative answer. It is true that when a
child more or less successfully reproduces a sound which falls
upon its ear, a like sound stimulus is afforded which may by a
circular process incite to renewed effort, and lead to yet more
successful reproduction. But when Professor Baldwin’s child,
between nine and ten months old, imitated certain movements
of the lips, there was no reproduction of the initiating visual
stimulus, A chick seeing its companions run away or crouch

IMITATION 183

will follow suit ; and this would commonly be termed an
imitative action; but there is here no reproduction of the
initiating stimulus. Very much of the behaviour which is
usually ascribed to imitation produces effects in consciousness
quite different from that of the suggestive stimulation. It is
only by selecting one’s examples that one finds in them evidence
in favour of Professor Baldwin’s “ circular process.”

Since, therefore, this circular mode of activity is neither a
characteristic of all conscious imitation, nor a distinguishing
mark of all adaptive organic action, the grounds on which
Professor Baldwin bases his extended usage of the term appear
to be fallacious. And in this usage we cannot follow him.

Turning now to Professor Thorndike’s very different con-
tention—that animals even so high as the cat and dog do not
imitate in the sense of forming an association leading to an
act from having seen another animal perform the act in a
certain way—we may first describe some of his ingenious
experiments designed to submit the matter to the test of ob-
servation under controlled conditions.”

Experiments were made with chicks in several ways. They
were, for example, placed in pens, from which, in each case,
“there was only one possible way of escape, to see if they
would learn it more quickly when another chick did the thing
several times before their eyes. The method was to give some
chicks their first trial with an imitation possibility, and their
second without, while others were given their first trial with-
out and their second with. If the ratio of the average time of
the first trial to the average time of the second is smaller in
the first class than it is in the second class, we may find
evidence of this sort of influence by imitation. Though
imitation may not be able to make an animal do what he
would otherwise not do, it may make him do quicker a thing
he would have done sooner or later anyway. As a fact, the
ratio is much longer. This is due to the fact that a chick,
when in a pen with another chick, is not afflicted by the dis- .
comfort of loneliness, and so does not try to get out. So the

* “ Animal Intelligence,” pp. 47-64.

184 SOCIAL BEHAVIOUR

other chick, who is continually being put in with him to teach

him the way out, really prolongs his stay in. This factor |

destroys the value of these quantitative experiments, and I do
not,” says Mr. Thorndike, “insist upon them as evidence
against imitation, though they certainly offer none for it.”

Chicks, from sixteen to thirty days old, were also placed in
boxes from which escape was open to them by such acts as
pecking at the door, stepping on a platform, or pecking at a
tack. The method of experiment was to put a chick in, leave
him from sixty to eighty seconds, then put in another who
knew the act, and on his performing it to let both escape. No
cases were counted unless the imitator apparently saw the
other do the thing. After about every ten such chances to
learn the act, the imitator was left in alone for ten minutes.
Out of thirteen cases tabulated only once was the act per-
formed, in spite of the ample chance for imitation. ‘I have
no hesitation,” adds Mr. Thorndike, “‘in declaring this one’s
act in stepping on the platform the result of mere accident,
and am sure that any one who had watched the experiments
would agree.”

To test the influence, if any, of imitation in cats, the
following method was adopted. A box was arranged with two
compartments separated by a wire screen. ‘‘ The larger of
these had a front of wooden bars with a door which fell open
when a string stretched across the top was bitten or clawed
down. The smaller was closed by boards on three sides and
by the wire screen on the fourth. Through the screen a cat
within could see the one to be imitated pull the string, go out
through the door thus opened, and eat the fish outside. When
put in this compartment, the top being covered by a large box,
a cat soon gave up efforts to claw through the screen, quieted
down, and watched more or less the proceedings going on in the
other compartment. Thus this apparatus could be used to test
the power of imitation. A cat who had no experience with
the means of escape from the large compartment was put in
the closed one; another cat, who would do it readily, was
allowed to go through the performance of pulling the string,

IMITATION 185

going out, and eating the fish. Record was made of the
number of times he did so, and of the number of times the
imitator had his eyes clearly fixed on him. . . . After
the imitatee had done the thing a number of times, the other
was put in the big compartment alone, and the time it took
him before pulling the string was noted and his general
behaviour closely observed. If he failed in five or ten or
fifteen minutes to do so, he was released and not fed. This
entire experiment was repeated a number of times. From the
times taken by the imitator to escape and from observation
of the way that be did it, we can decide whether imitation
played any part. . . . No one, I am sure, who had seen the
behaviour of the cats would have claimed that their conduct
was at all influenced by what they had seen. When they did
hit the string the act looked just like the accidental success of
the ordinary association experiment. But, hesides these personal
observations, we have in the impersonal time-records sufficient
proofs of the absence of imitation.” Some observations on
dogs are also described. From these it appears that the three
individuals on which experiments were made failed to learn
the way of getting out of a cage from seeing another dog
escape. One of them was also allowed to see another dog beg
for meat 110 times. But he never tried to imitate him and
thus secure a piece of meat asa reward. It therefore “seems
sure,” says Mr. Thorndike, “that we should give up imitation
as an a priori explanation of any novel intelligent performance.
To say that a dog who opens a gate, for instance, need not
have reasoned it out if he had seen another dog do the same
thing, is to offer instead of one false explanation another
equally false. Imitation in any form is too doubtful a factor
to be presupposed without evidence.”

Professor Thorndike is of opinion that monkeys are pro-
bably imitative in ways beyond the capacity of dogs and cats ;
but, at the time of writing, he had not substantiated his opinion,
by analogous experiments. If so, it will perhaps prove that
they are rational beings in the narrower sense defined in
a previous chapter of this work. For it appears that the

186 SOCIAL BEHAVIOUR

kind of imitation which Mr. Thorndike’s experiments go far
to disprove, is what we may term reflective imitation. A
cat with no experience of the means of escape, one that has
tried to get out of the box by chance efforts in many directions
and has failed, sees another cat perform an act acquired in
this way, and learns nothing from the sight. This, no doubt,
proves that the cat had not in any sense grasped the nature of
the problem before it, had no notion of just where the difficulty
lay, had not the wit to see that the performance of the other
cat supplied the missing links in its own previous behaviour.
It is questionable whether such missing links could be supplied
in this way in the absence of some powers of reflection. The
cat is unable to form an association, leading to an appropriate
act, from having seen another animal perform the act in a
certain way, partly because it cannot perceive the reason of its
previous failure, and see that the other’s performance affords
the requisite clue. The whole gist of the chance experience
interpretation of animal behaviour is that there must be
chance experience to build on. The cat cannot gain this by
looking on never so intently, unless it be provided with a
rational as well as a sensory eye. The act of pulling the
string has been reached by the successful cat through the
gradual elimination of many failures; it is a differentiated
act, having no place in the previous experience of the kitten.
It has never entered into the conscious situation, and cannot
be supplied at will by a non-rational being.

As Mr. Thorndike himself says, “no cat can form an
association leading to an act unless there is included in the
association an impulse of its own which leads to the act.” *
By “impulse,” Mr. Thorndike ‘‘ means the consciousness
accompanying a muscular innervation apart from that feeling
of the act which comes from seeing one’s self move, from feel-
ing one’s body in a different position, etc. It is the direct
feeling of doing as distinguished from the idea of the act
done gained through eye, etc. . . The act in this respect
of being felt as to be done or as doing is in animals the

* Op. cit., pp. 66, 14, 15.

IMITATION 187

important thing, is the thing which gets associated, while the
act as done, as viewed from outside, is a secondary affair.”
I take it that by “impulse” is here meant what Dr. Stout
would term the direct experience involved in conation.* If
it have a place in experience distinguishable from that of
stimulation and response it is included in what I have on a
former page spoken of as the consciousness of behaviour as
such, which was said to be essential. And I am surprised
that Mr. Thorndike should have supposed that I believe that
this could by any animal be “supplied at will.’ In any case
it seems probable, as the result of observation, that unless the
consciousness of behaving in a specific manner has entered
into the situation as developed in experience it cannot in
animals enter into any subsequent representative complex.
And it is the absence of such consciousness of behaving in a
specific manner which the sight of the escaping cat fails to
supply in Mr. Thorndike’s experiments.

Interesting and valuable as these experiments are, they are
open to the criticism to which, as we have seen, his other
experiments are also open—that the conditions are abnormal
and cramped. Apart from reflective imitation, which they
tend to disprove, they do not conduce to the kind of conscious
situation which appears to be most favourable for the develop-
ment of intelligent imitation founded on hereditary tenden-
cies and propensities. It is through such imitation that, as
Herr Groos says,t “animals learn perfectly those things for
which they have imperfect hereditary dispositions.” The
kind of situation which conduces to such intelligent imitation
is that which involves the attitude of attention and interest
rising, when these are sufficiently varied in their direction, into
what is spoken of as curiosity. These, in their natural occur-
rence in animals, are parts of, or in any case accompaniments
of the conative attitude—they are connected with activities
and impulsive tendencies to behaviour. If attention and
interest are directed to the behaviour of another animal, the

* Cf. infra, p. 235.
+ “The Play of Animals,” Eng. trans., p. 79.

138 SOCIAL BEHAVIOUR

conative attitude is that of imitation. Miss Romanes has
described how skilfully a capuchin imitated the actions neces-
sary to unlock a trunk. Jt does not seem necessary to assume
that reflective imitation is here exemplified. The monkey
need not regard the key and lock as the related parts of a
puzzle to be practically solved, need not have any free idea of
the difficulty it presents, need not in unlocking the trunk
grasp the true nature of the difficulty or have any conception
of its solution. Every several act of the capuchin, the seizing
the key, the directing it here or there, and so on, is already
supplied with the impulse of which Dr. Thorndike speaks.
Attention, itself charged with impulse, directs and combines
these pre-existing impulses to a new end. And since that
which directs the attention is the act of another, we call the
procedure imitative. But the varied and persistent effort
differs in no essential respect from that of a two days’ chick,
which pecks again and again at some speck which catches its
eye, or that of a nestling jay, which will peck for long at some
rail or piece of wire in its cage, twisting and turning its bill
in many and varied ways. And success in opening the trunk
is reached by the capuchin, not, it would seem, through any
real appreciation of the essential kernel of the practical pro- :
blem, but through the chance results of many varied efforts.
Although in no other animals is it developed to so high a
degree as in the monkeys, interest in the doings of others is
an attitude by no means rare, and affords the basis of intelli-
gent imitation. Perhaps the conditions in Dr. Thorndike’s
experiments were not the best for the development of such
interest in the procedure of another. And in any case the
imitation of a particular mode of procedure, reached by the
gradual defining of the impulse, could hardly be expected in
the absence of the series of experiences by which that defini-
tion had been reached, unless the cat were capable of what
has been above spoken of as reflective imitation.

If, then, we agree to exclude from the category of imita-
tive behaviour in animals, on the one hand, any “ circular
process”? which may occur in the same individual, and on the

IMITATION 189

other hand any reflective imitation, such as is so important a
factor in human education, it remains to be seen what may be
fairly included in this category.

It is probable that in animals imitation has its foundations
in instinctive behaviour, of which it may be regarded as the
characteristically social type. If one of a group of chicks
learn by casual experience to drink from a tin of water, others
will run up and peck at the water, and thus learn to drink.
A hen teaches her little ones to pick up grain or other food
by pecking on the ground and dropping suitable materials
before them, while they seemingly imitate her action in seizing
the grain. One may make chicks and pheasants peck by
simulating the action of a hen with a pencil point or pair of
fine forceps. According to Mr. Peal, the Assamese find that
young jungle pheasants will perish if their pecking responses
are not thus stimulated ; and Professor Claypole tells me that
this is also the case with young ostriches hatched in an
incubator. A little pheasant and guinea-fowl followed two
older ducklings, one wild, the other tame, and seemed to wait
upon their bills, to peck when they pecked, and to be guided
by their actions. It is certainly much easier to bring up
young birds if older birds are setting an example of eating
and drinking ; and instinctive acts, such as scratching the
ground, are performed earlier if imitation be not excluded.
If a group of chicks have learnt to avoid cinnabar cater-
pillars, and if then two or three from another group are
introduced and begin to pick up the caterpillars, the others
will sometimes again seize them, though they would otherwise
have left them untouched. One of my chicks, coming upon
a dead bee, gave the danger or alarm note; another at some
little distance at once made the same sound. A number of
similar cases might be given ; but what impresses the observer
as he watches the early development of a brood of young
birds, is the presence of an imitative tendency which is
exemplified in many little ways not easy to describe in detail.
It is probable, however, that these imitative tendencies or
propensities are not wholly indefinite. The young birds do

190 SOCIAL BEHAVIOUR

not imitate any actions, but behaviour of certain specific types,
the imitation of which has been engrained through the action
of natural selection.

What generalization, then, can be drawn from this some-
what indefinite group of facts, to which many others of like
import could be added from observations on the young of
mammals? What is their relation to instinctive procedure in
general? It would seem that they are characterized by a
special relation of the external stimulus to the response.
When this stimulus is afforded by the behaviour of another
animal, and the responsive behaviour it initiates is similar to
that which affords the stimulus, such behaviour may be termed
imitative. A chick sounds the danger note; this is the
stimulus under which another chick sounds a similar note,
and we say that the one imitates the other. Such an action
may be described as imitative in its effects, but not imitative
in its purpose. Only from the observer’s standpoint does such
instinctive behaviour differ from other modes of congenital
procedure. It may be termed biological, but not psychological,
imitation. And if it be held that the essence of imitation lies
in the purpose so to imitate, we must find some other term
under which to describe the facts. This does not seem neces-
sary, however, if we are careful to qualify the term “ imita-
tion” by the adjective “instinctive” or “biological.” And
the retention of the term serves to indicate that this is the
stock on which deliberate imitation is eventually grafted.

The fact that instinctive imitation leads, under natural
conditions, to behaviour which is already familiar to us
in the species concerned, prevents us from recognizing the
influence of this social factor so easily as might otherwise be
the case. The abnormal arrests our attention more readily
than the normal, and hence the cases commonly cited are
generally those which strike us as unusual, such as the
imitation of human sounds by the parrot. But if the
young inherit a tendency to imitate certain actions of their
parents, and if there is among the members of a gregarious
species such instinctive imitation as shall tend to keep them

IMITATION 191

gregarious, we have here a social factor in animal life of no
slight importance. Just as the higher type of reflective
imitation is of great value in bringing the human child to
the level of the adults who form the family and social environ-
ment, so, too, does the sub-conscious instinctive imitation of
the lower animals bring the young bird or other creature into
line with the members of its own species. In broods of chicks
brought up under experimental conditions, there are often one
or two more active, vigorous, intelligent, and mischievous
birds. These are the leaders of the brood; the others are
their imitators. Their presence raises the general level of
intelligent activity. Remove them, and the others show a less
active, less inquisitive, less adventurous life. They seem to
lack initiative. From which one may infer that imitation
affords to some extent a means of levelling up the less intelli-
gent to the standard of the more intelligent ; and of supply-
ing a stimulus to the development of habits which would
otherwise be lacking. When a mongrel pup, whose develop-
ment Dr. Wesley Mills watched and has described, was intro-
duced to the society of other dogs, its progress was, he tells
us, ‘ extraordinarily rapid.”

Instinctive imitation thus introduces into the conscious
situation certain modes of behaviour, and if the development
of the situation as a whole is pleasurable, there will be a ten-
dency to its redevelopment, under the guidance of intelli-
gence, on subsequent occasions. As in the case of other
instincts and propensities, there is given through inheritance
a more or less definite outline sketch of social procedure,
which intelligence further defines, and refines, and shapes to
more delicate issues. As a rule, however, intelligence does
not tend to make the imitation as such more perfect. It may
perfect the behaviour, but not necessarily on imitative lines.
In the case, however, of the song and call-notes of birds, and
not improbably the sounds of other animals, there does seem
a predisposition to render the imitation as such more perfect.
The facts, as afforded by such birds as the magpie, jay,
starling, marsh-warbler, and mocking-bird, are familiar ; and

- 192 SOCIAL BEHAVIOUR

I have elsewhere * given some account of them. It may be
specially noted that we have in this case that circular mode
of activity on which, as we have seen, Professor Mark Bald-
win lays so much stress. Professor Thorndike seems to regard
the phenomena presented by imitative birds as somewhat of a
mystery, and as the result of a specialization removed from
the general course of mental development. And he says that,
until we know whether there is in birds which repeat sounds
any tendency to imitate in other lines, we cannot connect
these phenomena with anything found in the mammals, or
use them to advantage in a discussion of animal imitation as
the forerunner of human. Upon the view, however, that such
imitation is primarily instinctive and only secondarily intelli-
gent, there seems no reason why we should expect to find
imitation in birds running along any other lines than those
which the hereditary instinct has marked out. And so far
from being unable to use the phenomena to advantage in a
discussion of animal imitation as a forerunner of human, we
may perhaps see in them the best examples, other than those
afforded by apes, of that intelligent imitation which is the
precursor of the rational and reflective imitation of the boy or
girl.

In the case of the human child we may see the three stages
in the development of imitation. First, the instinctive stage,
where the sound which falls upon the ear is a stimulus to the
motor-mechanism of sound production. Secondly, the in-
telligent stage of the profiting by chance experience. Intelli-
gence, as we have seen, aims at the reinstatement of pleasurable
situations, and the suppression of those which are the reverse.
The sound-stimulus, the motor effects in behaviour, and the
resulting sound-production coalesce into a conscious situation,
which appears to be pleasurable or the reverse, according as
the sound produced resembles or not the initiating sound-
stimulus. If we assume that the resemblance of the sounds he
utters to the sounds he hears is itself a source of pleasurable
satisfaction (and this certainly seems to be the case), intelligence,

* © Habit and Instinct,” pp. 174-180,

INTERCOMMUNICATION 193

without the aid of any higher faculty, will secure accommoda-
tion and render imitation more and more perfect. And this
appears to be the stage reached by the mocking-bird or the
parrot. But the child soon goes further. He reflects upon
the results he has reached ; he at first dimly, and then more
clearly realizes that they are imitative ; and his later efforts at
imitation are no longer subject to the chance occurrence of
happy results, but are based on a scheme of behaviour which is
taking form in his mind, are deliberate and intentional, and
are directed to a special end more or less clearly perceived as
such. He no longer imitates like a parrot; he begins to
imitate like a man, and may, by the study of good models and
the maintenance of a high ideal, acquire the moving cadences
of an orator.

According to our interpretation, instinctive imitation is a
factor of wide importance in animal behaviour, intelligent
imitation, arising in close connection with interest in the
doings of others, is a co-operating factor, but of intentional and
reflective imitation there is at present no satisfactory evidence
in any animal below man.

II.—IntTERCOMMUNICATION

The foundations of intercommunication, like those of
imitation, are laid in certain instinctive modes of response,
which are stimulated by the acts of other animals of the same
social group. These have been fostered by natural selection as
a means of social linkage furthering the preservation, both of
the individual and of the group.

Some account has already been given of the sounds made
by young birds, which seem to be instinctive and to afford an
index of the emotional state at the time of utterance. That in
many cases they serve to evoke a like emotional state and
correlated expressive behaviour in other birds of the same
brood cannot be questioned. The alarm note ofa chick
will place its companions on the alert; and the harsh
“krek ? of a young moor-hen, uttered in a peculiar crouching

te)

194 SOCIAL BEHAVIOUR

attitude, will often throw others into this attitude, though the
maker of the warning sound may be invisible. That the
cries of her brood influence the conduct of the hen is a
matter of familiar observation; and that her danger signal
causes them at once to crouch or run to her for protection
is not less familiar. No one who has watched a cat with
her kittens, or a sheep with her lambs, can doubt that such
“dumb animals” are influenced in their behaviour by sug-
gestive sounds. The important questions are, how they
originate, what is their value, and how far such intercom-
munication—if such we may call] it—extends.

There can be but little question that in all cases of animals
under natural conditions such behaviour has an instinctive
basis. Though the effect may be to establish a means of com-
munication, such is not their conscious purpose at the outset.
They are presumably congenital and hereditary modes of
emotional expression which serve to evoke responsive behaviour
in another animal—the reciprocal action being generally in
its primary origin between mate and mate, between parent
and offspring, or between members of the same family group.
And it is this reciprocal action which constitutes it a factor
in social evolution. Its chief interest in connection with
the subject of behaviour lies in the fact that it shows the
instinctive foundations on which intelligent and eventually
rational modes of intercommunication are built up. For
instinctive as the sounds are at the outset, by entering into
the conscious situation and taking their part in the association-
complex of experience, they become factors in the social life
as modified and directed by intelligence. To their original
instinctive value as the outcome of stimuli, and as themselves
affording stimuli to responsive behaviour, is added a value for
consciousness in so far as they enter into those guiding situa-
tions by which intelligent behaviour is determined. And if
they also serve to evoke, in the reciprocating members of the
social group, similar or allied emotional states, there is thus
added a further social bond, inasmuch as there are thus laid
the foundations of sympathy.

INTERCOMMUNICATION 195

‘What makes the old sow grunt and the piggies sing and
whine ?” said a little girl to a portly substantial farmer. “I
suppose they does it for company, my dear,” was the simple
and cautious reply. So far as appearances went, that farmer
looked as guiltless of theories as man could be. And yet he
gave terse expression to what may perhaps be regarded as the
most satisfactory hypothesis as to the primary purpose of
animal sounds. They are a means by which each indicates to
others the fact of his comforting presence; and they still, to
a large extent, retain their primary function. The chirping
of grasshoppers, the song of the cicada, the piping of frogs in
the pool, the bleating of lambs at the hour of dusk, the lowing
of contented cattle, the call-notes of the migrating host of
birds—all these, whatever else they may be, are the reassuring
social links of sound, the grateful signs of kindred presence.
Arising thus in close relation to the primitive feelings of social
sympathy, they would naturally be called into play with special
force and suggestiveness at times of strong emotional excite-
ment, and the earliest differentiations would, we may well
believe, be determined along lines of emotional expression.
Thus would originate mating cries, male and female after
their kind; and parental cries more or less differentiated into
those of parent and offspring, the deeper note of the ewe
differing little save in pitch and timbre from the bleating of
her lamb, while the cluck of the hen differs widely from the
peeping note of the chick in down. Thus, too, would arise
the notes of anger and combat, of fear and distress, of alarm
and warning. If we call these the instinctive language of
emotional expression, we must remember that such “language”
differs markedly from the “language” of which the sentence
is the recognized unit.

It is, however, not improbable that, through association in
the conscious situation, sounds, having their origin in emotional
expression and evoking in others like emotional states, may
acquire a new value in suggesting, for example, the presence of
particular enemies. An example will best serve to indicate my
meaning. “In the early dawn of a grey morning,” says Mr.

196 SOCIAL BEHAVIOUR

H. B. Medlicott,* “I was geologizing along the base of the
Muhair Hills in South Behar, when all of a sudden there was
a stampede of many pigs from the fringe of the jungle, with
porcine shrieks of sawve-qui-peut significance. After a short
run in the open they took to the jungle again, and in a few
minutes there was another uproar, but different in sound and
in action; there was a rush, presumably of the fighting
members, to the spot where the row began, and after some
seconds a large leopard sprang from the midst of the scuffle.
In a few bounds he was in the open, and stood looking back,
licking his chaps. The pigs did not break cover, but con-
tinued on their way. They were returning to their lair after
a night’s feeding on the plain, several families having com-
bined for mutual protection ; while the beasts of prey were
evidently waiting for the occasion. I was alone, and, though
armed, I did not care to beat up the ground to see if in either
case a kill had been effected. The numerous herd covered a
considerable space, and the scrub was thick. The prompt
concerted action must in each case have been started by the
special cry. I imagine that the first assailant was a tiger, and
the case was at once known to be hopeless, the cry prompting
instant flight, while in the second case the cry was for defence.
It can scarcely be doubted that in the first case each adult pig
had a vision of a tiger, and in the second of a leopard or some
minor foe.”

Tf we accept Mr. Medlicott’s interpretation as in the main
correct, we have in this case: (1) common action in social
behaviour, (2) community of emotional state, and (3) the sug-
gestion of natural enemies not unfamiliar in the experience of
the herd. Under uniform conditions of experience the alarm-
notes of some birds may well call up, re-presentatively, salient
features in previous situations. Unquestionably, in the parrot,
the word-sounds they imitate become associated with definite
objects of sense-experience. In the following case, a particular

* «The Evolution of Mind in Man,” footnote, pp. 25, 26. Quoted in
“ Introduction to Comparative Psychology,” from which the comments on
it are extracted.

INTERCOMMUNICATION 197

sound appeared to be suggestive of a particular sense-idea in
the dog. The parent blackbirds, which built near a house in
Clifton, were wont to give the alarm-note when marauding
cats appeared in sight. This sound, it would seem, became
definitely associated, in the experience of a terrier, with the
animals the presence of which called it forth ; and on hearing
the alarm note the dog would rush out into the garden,
apparently, as I am informed by his mistress, in fullest ex-
pectation of a pleasant worry. It is a not improbable hypothesis,
therefore, that in the course of evolution the initial value
of uttered sounds is emotional; but that on this may be
grafted in further development the indication of particular
enemies. If, for example, the cry which prompts instant flight
among the pigs is called forth by a tiger, it is reasonable to
suppose that this cry would give rise to a representative generic
image of that animal having its influence on the conscious
situation. But if the second cry, for defence, was prompted
sometimes by a leopard and sometimes by some other minor
foe, then this cry would not give rise to a re-presentative image
of the same definiteness. Whether animals have the power of
intentionally differentiating the sounds they make to indicate
different objects, is extremely doubtful. Can a dog bark in
different tones to indicate “cat” or “rat,” as the case may
be? Probably not. It may, however, be asked why, if a
pig may squeak differently, and thus, perhaps, incidently
indicate on the one hand “tiger” and on the other hand
“leopard,” should not a dog bark differently, and thus indicate
appropriately “cat” or “rat”? Because it is assumed that the
two different cries in the pig are the instinctive expression
of two different emotional states, and Mr. Medlicott could
distinguish them; whereas, in the case of the dog, we can
distinonish no difference between his barking in the one
case and the other, nor do the emotional states appear to be
differentiated. Of course, there may be differences which we
have failed to detect. What may be regarded, however, as
improbable, is the intentional differentiation of sounds by
barking in different tones with the purpose of indicating “ cat.”

198 SOCIAL BEHAVIOUR

or “rat.” Mr. R. L. Garner, in a work * which unfortunately
contains much hasty and immature generalization, distinguished
nine sounds made by capuchins. But none of these, so far as
can be gathered from the data given, is necessarily indicative
of a particular object. All of them may be emotional ex-
pressions of satisfaction, discontent, alarm, apprehension, and
so forth. In any case, there is no evidence for that inten-
tional employment of sounds, to the realized end of inter-
communication, which would involve the exercise of an
incipient rational faculty. Such powers of intercommuni-
cation as animals possess are based on direct association,
and refer to the here and the now. A dog may be able to
suggest to his companion the fact that he has descried a
worriable cat; but can a dog tell his neighbour of the
delightful worry he enjoyed the day before yesterday in the
garden where the man with the biscuit-tin lives? Probably
not, bark he never so expressively.

Although some anecdotes are commonly interpreted as
affording evidence of descriptive intercommunication among
animals, we need the decisive results of experiment before this
view can be unreservedly accepted. Sir John Lubbock, now
Lord Avebury, made careful experiments with ants, and dis-
cusses the question with his customary lucidity and impar-
tiality. ‘Much of what has been said,” he writes,f ‘as to the
powers of communication possessed by bees and ants depends
on the fact that if one of them in the course of her rambles
has discovered a supply of food, a number of others soon find
their way to the store. This, however, does not necessarily
imply any power of describing localities. If the ants merely
follow a more fortunate companion, or if they hunt her by
scent, the matter is comparatively simple ; if, on the contrary,
the others have the route described to them, the case becomes
very different.” Experiments were therefore made to decide
the question. For example, when an ant returned from the
discovered store of food to the nest, and then emerged with a

* «The Speech of Monkeys.”
t “Scientific Lectures,” pp. 112, 118.

INTERCOMMUNICATION 199

following of other ants, she was taken up on a slip of paper
and transferred to the food. The followers, thus deprived of
their leader, in nearly all cases failed to find the store. “I
conclude, then,” says Lord Avebury, “that when large
numbers of ants come to food they follow one another, being
also, to a large extent, guided by scent. The fact, therefore,
does not imply any considerable power of intercommunication.”
There are, moreover, some circumstances which seem to
strengthen this conclusion. For instance, “if a number of
slave-ants are put in a box, and if in one corner a dark place
of retreat be provided for them, with some earth, one soon
finds her way to it. She then comes out again, and going up
to one of the others, takes her by the jaws and carries her
to the place of shelter. They then both repeat the same
manceuvre with other ants, and so on until all their com-
panions are collected together. Now, it seems difficult to
imagine that so slow a course would be adopted, if they
possessed any power of communicating description.”

Lord Avebury is, however, of opinion that such insects
can transmit simpler ideas. He found, for example, that
where ants were put to a large and a small store of larve
under similar circumstances, a greater number of insects
followed the ant that had discovered the larger store. This
may, indeed, have been due rather to a difference in manner
than to any intentional communication ; but the fact remains
that through some difference of behaviour there resulted
suggestive effects on other members of the community.

But although there can be little doubt that the behaviour
of social insects has suggestive value for others, it may still be
regarded as very doubtful whether they are able to communi-
cate information to one another by any system of language’or
signs, purposively employed as a system to this end. The
distinguished geologist, Hague, communicated to Darwin * the
effects on ants of crushing some of their number as they pro-
ceeded along a definite trail. “As soon as those ants which
were approaching arrived near to where their fellows lay dead

* Nature, vol. vii., p. 443.

200 SOCIAL BEHAVIOUR |

”

and suffering, they turned and fled with all possible haste.
“ When such an ant, returning in fright, met another approach-
ing, the two would always communicate, but each would
pursue its own way, the second ant continuing its journey to
the spot where the first had turned about, and then following
that example.” There seems nothing to show that the
“communication ” here was effective.

From the many anecdotes of dogs calling others to their
assistance, or bringing others to those who feed them or treat
them kindly, we may indeed infer the existence of a social
tendency and of the suggestive effects of behaviour, but we
cannot derive conclusive evidence of anything like descriptive
communication. And although domestic animals may learn
or be taught to associate the words we utter with certain acts
or things, or may even, in a sense, communicate their wishes
to us by special modes of behaviour—as in the case of Lord
Avebury’s poodle, Van,* who was taught to bring cards on which
such words as ‘“‘ Food” or “ Out” were printed, and in that of
a eat which touches the handle of the door when she wants
it opened for her,—still, all these are founded on direct
association, and are in a line with the act of Mr. Thorndike’s
cat, which licked herself or scratched herself when imprisoned
in a cage, such act having entered into the association-complex.

Such intentional communication as is to be found in animals,
if indeed we may properly so call it, seems to arise by an
association of the performance of some act in a conscious
situation involving further behaviour for its complete develop-
ment. Thus the cat which touches the handle of the door
when it wishes to leave the room has had experience in which
the performance of this act has coalesced with a specific
development of the conscious situation. The case is similar
when your dog drops a hall or stick at your feet, wishing you
to throw it for him to fetch. And on these lines may probably
be interpreted such behaviour as Romanest thus described :—
“ Terrier A being asleep in my house, and terrier B lying on a

* « The Senses of Animals,” p. 277.
t “Mental Evolution in Man,” p. 100.

INTERCOMMUNICATION 201

wall outside, a strange dog, C, ran along below the wall on the
public road, following a dog-cart. Immediately on seeing C,
B jumped off the wall, ran upstairs to where A was asleep,
woke him up by poking him with his nose in a determined
and suggestive manner, which A at once understood as a sign:
he jumped over the wall and pursued the dog OC, although C
was by that time far out of sight round a bend in the road.”
Romanes did not probably intend to imply that A by poking
B, conveyed specific information that there was another dog,
C, which had proceeded in a particular direction. That would
be descriptive communication. The meaning attaching to A’s
action was presumably similar to that which characterizes
other “ meaning ” for intelligent animals—the development of
the situation on lines marked out by previous experience.
Still, it is clear that such an act would be the perceptual
precursor of the deliberate conduct of the rational being by
whom the sign is definitely realized as a sign, the intentional
meaning of which is distinctly present to thought. This
involves a judgment concerning the sign as an object of
thought ; and this is probably beyond the capacity of the dog.
For, as Romanes himself says,* “it is because the human
mind is able, so to speak, to stand outside of itself, and thus
to constitute its own ideas the subject-matter of its own
thought, that it is capable of judgment, whether in the act of
conception or in that of predication. We have no evidence
to show that any animal is capable of objectifying its own
ideas ; and, therefore, we have no evidence that any animal is
capable of judgment.”

It seems, therefore, that the sounds made by animals, and
certain other modes of behaviour, may be regarded as primarily
instinctive acts which have been evolved with the biological
end of affording suggestive stimuli furthering intercom-
munication between the members of the social group. Their
performance, however, affords data to consciousness, which
intelligence makes use of in the guidance of behaviour in
accordance with the results of experience. And since the

* Op. cit. p. 175. :

202 SOCIAL BEHAVIOUR

similar acts performed by the socially linked members are in
many cases closely connected with emotional states, there arises
the further social link of community of feeling—that which,
perhaps, more than anything else conduces to community of
action and similarity of social behaviour. Occasionally
particular sounds or special acts may, through constant and
uniform association, indicate particular objects, such as natural
enemies. But there does not appear to be convincing evidence:
of any intentional differentiation of the means of communica-
tion, or of any use of sounds for descriptive ends.

Still, just as the instinctive imitation we considered in the
last section may be regarded as the precursor, in the animal
world, of the reflective and rational imitation of which we may
watch the development in children, so may instinctive modes
of intercommunication be regarded as supplying the foundations
on which deliberate and intentional communication may be
based. And here imitation will be a co-operating factor. We
see in the early stages of the development of children’s language
how large a share simple and direct association takes in the
process. For a while, indeed, there seems to be this and nothing
more. But gradually there arises a realization of a further
import and purpose in the hitherto isolated associations. It is
seen that they symbolize elements in that incipiently rational
scheme of thought and things which is beginning to take form,
in the child’s mind. The relationships which hold good within
the conscious situations of daily life begin to occupy the focus
of attention, and hitherto unappreciated word-sounds are per-
ceived to stand out as signs for these relationships. Of course
the relationships * are implicit in the conscious situations of
the higher animals and of infants. Only by reflection can
they become explicit, and rivet the attention. Something is
needed to bring them into prominence and focus the mental
eye upon them. And descriptive intercommunication supplies
this need. If a description, even the simplest, is to be
apprehended or presented to the apprehension of others, then

* Compare chap. xiii, on “The Perception of Relations,” in my
“Introduction to Comparative Psychology.”

INTERCOMMUNICATION 203

the relationships must be rendered explicit. Try to describe
an ordinary visual scene, or the most commonplace sequence
of events, and see if you can do so without making clear to the
mind the relationships involved. The thing is impossible. An
infant or a dog cannot understand the simplest possible
description, because the words and suffixes which indicate the
relationships have no meaning. The words which stand for
substantive impressions may have suggestive value through
direct association. The word “cat” or “rats” may have for
the dog a very definite suggestive value; and hence some
people fancy that when they say to their dog, “ There is a cat
in the garden,” the animal understands what they say. But it
is quite sufficient to suppose that the word “cat ”’ has suggestive
force, all the rest being for the dog mere surplusage of sound.
When we talk to our four-footed companions, how much can
they be said to understand of what we say ? Perhaps a score
of words have for a dog a definitely suggestive value, each
associated with some simple object or action. ‘ Out,” “down,”
“up,” “walk,” “biscuit,” “cat,” “fetch,” and so forth elicit
appropriate responses. Even with these, tone is more sug-
gestive than articulation, and in each word the salient feature
is the chief guide. When I said “ Whisky,” for example, to
my fox-terrier, he would at once sit up and beg; not because
his tastes were as depraved as those of his master, but because
the ask sound, common both to “ whisky” and “ biscuit,” was
what had for his ears the suggestive value.

In a paper on the “Speech of Children,” * Mr. 8. 8.
Buckman exhibits the animal stage in the incipient speech of
the human infant. We cannot here discuss, still less criticize,
his paper. One or two examples will serve to illustrate how
instinctive sounds may serve as the basis for subsequent
speech. He regards ma as primarily a forcible expression of
an emotional state. “If the child require attention it makes
the loudest noise which it can produce; the parting of the
lips and opening of the mouth to the widest extent while the
full volume of breath is emitted produces the sound ma.” At

* Nineteenth Century, May, 1897, pp. 793-807.

204 SOCIAL BEHAVIOUR

first the sound seems to have the value of a simple expression
of an emotional state. “ But if the infant require attention it
is its mother whom it wants, and from whom it receives this
attention ; therefore ma very soon comes to be recognized as
the call for mother, and, by a further step in development, as
the name for mother.” Here, if we accept the interpretation,
we have the passage from the emission of a sound as the
expression of emotion to the use of the sound from its associa-
tion with a particular object of sense-experience to indicate
that object. Similarly, according to Mr. Buckman, with hah.
At first “a strong sign of displeasure at anything nasty to the
taste,” it passes, we are told, into a symbol for the bad ; hence
xaxos; and is perhaps narrowed down to the particularly
offensive xdxxy. Da and ta are regarded as recognition sounds,
the former being associated eventually with the father, the
latter with strangers. This appears somewhat hypothetical,
but, granting the accuracy of Mr. Buckman’s interpretation,
these sounds also illustrate again the transition from the
expression of an emotion to sounds indicative of particular
objects of experience.

Interesting, however, as are such observations on the animal
stage of sound-production in the human infant, they do not
touch the crucial period in the development of language. Mr.
Buckman, indeed, regards as a remarkably dogmatic assertion
Professor Max Miiller’s dictum that “the one great barrier
between the brute and man is language; ” and he tells us that
“there are more than twelve different words in the language
of fowls,” on which assertion, in turn, the distinguished linguist
whom he criticizes might have something piquant to say. No
doubt the difference of opinion turns on the definition of the
word “language.” But if, as is now generally accepted, the
sentence and not the word is the distinguishing unit in
language, and the copula in some form, explicit or implicit, is
the pivot of the sentence, the wisest hen is probably incapable
of language. The word becomes an element in language—a
word proper—only when it assumes the office of a part of
speech, that is to say, a constituent element in an interrelated

SOCIAL COMMUNITIES OF BEES AND ANTS 205

whole. The animal “word,” if we like so to term it, is an
isolated brick; a dozen, or even a couple of hundred such
bricks do not constitute a building. Language, properly so
called, is the builded structure, be it a palace or only a cottage ;
hen language, or monkey language, is, at best, so far as we at
present have evidence, an unfashioned heap of bricks. It is
just because language is the expression of a portion of a scheme
of thought that it indicates in the speaker the possession of a
rational soul, capable of perceiving and symbolizing the
relationships of things as reflected in thought.

Herein lies the practical value, for human advance in
mental development, of language as a means of descriptive —
intercommunication. It renders explicit relationships other-
wise merely implicit, and forces them to the front ; and since
these relationships are the stuff of which knowledge is built —
without the realization of which any complex ideal scheme is
impossible of attainment—the importance of descriptive inter-
communication can scarcely be overestimated. And though
there is no conclusive evidence of its occurrence among
animals, yet we have in them the instinctive and intelligent

basis on which in due course of evolution it may be securely
based.

III.—Socran ComMMUNITIES OF BEES AND ANTS

Apart from human societies the most noteworthy social
communities of animals are found among insects, especially in
ants, bees, wasps, and termites. It is true that in the mam-
malia we find such communities as the troop of apes, the herd
of cattle, the pack of wolves, the school of porpoises, the so-
called “rookeries” of seals, and the colonies of “ prairie
dogs” and of beavers; and that among birds there are
analogous communities. Undoubtedly the temporary or per-
manent association of many individuals is in such cases an
advantage to the race, and confers mutual benefits on the
associates. But in none of these cases is division of labour
carried to such a high degree as among the social insects.

206 SOCIAL BEHAVIOUR.

And it is through such division of labour that the social com-
munity reaches its highest expression.

It is a somewhat remarkable fact that in man, where we
find the social division of labour brought to a high pitch of
perfection, and carried out with great nicety of accommoda-
tion to those circumstances which civilization has rendered
extremely complicated, there is no organic differentiation of
structure among the co-operating individuals; whereas, so
low down in the scale of life as the colonial polype, Hydrac-
tinta, which is often found growing on the shells oceupied by
hermit crabs, there are at least three kinds of differentiated
individuals: nutritive polypes with mouth and tentacles ;
mouthless sensitive members ; and others whose sole office is
reproduction. But these differentiated individuals in the
colonial zoophytes are connected at their bases by a common
flesh ; and the division of labour is a product of organic
evolution, and is probably not in any degree determined or
guided by consciousness. We may say, then, that the division
of labour in the zoophyte is wholly physical, whereas in man
it is chiefly conscious or psychical; as is also the bond of
union between the several members of the colony. Inter-
mediate between these extremes stand the social insects. In
them there is no physical bond of union, for each individual
is distinct and separate ; the social linkage is in some degree
conscious under the conditions of their nurture; and the
division of labour is partly conscious, though probably in
large degree based on instinctive foundations, and partly the
outcome of an organic differentiation of structure seen in the
reproductive members’and in the sterile workers, as exempli-
fied in the common wood ant (Fig. 24). In some cases the
workers themselves may be divided into different castes.

So much has been written—and well written—on the
social life of insect communities, that it will here suffice to
indicate some of the problems which arise when we endeavour
to interpret the modes of behaviour which have been carefully
observed. In the honey-bee we have the well-known differen-
tiation of structure into drones or effective males, queens

SOCIAL COMMUNITIES OF BEES AND ANTS 207

or egg-laying females, and workers or ineffective females, in
which the development of the reproductive organs is arrested
or modified. Distinct modes of behaviour are correlated with
these structural differences. When a swarm of bees leaves a
hive it generally consists of the old queen-mother and a
certain number of the workers which are her offspring. When
they have found new quarters, or have been safely housed
under. domestication, the workers busy themselves in making
the cells in which the queen may lay her eggs, and in which
food may be stored. In doing this the bees act in concert,

Fic. 24.—Wood ant. 1, Queen; 2, male; 3, worker (from Shipley).

and though the mathematical accuracy of the form and size
of the cells has been much exaggerated, the comb which
results is a very beautiful and well-adapted product of mutual
co-operation in joint labour. And though intelligence may,
under special circumstances, modify the method of procedure
there can be little ‘doubt that comb-building is primarily due
to inherited instinct. The cells are not, however, all of the
same size, those for the drones being somewhat larger than
the cells in which the workers are reared, while much larger
and differently shaped cells are prepared for the future queens.
If instinctive therefore—as it seems to be in the main—the
behaviour runs into different lines, the immediate causes of
which, internal or external, we are not able accurately to
assign.

The reproductive behaviour of egg-laying in the queen-
mother is also instinctive. It is believed that the drones are
developed from eggs from which the queen bee withholds the
fertilizing fluid, which she retains for months or years after the

208 SOCIAL BEHAVIOUR

nuptial flight, stored in a special receptacle. And the size and
shape of the drone-cell may supply the stimulus through which
her behaviour in this respect is determined. But she lays
similarly fertilized eggs in both the worker-cells and the
queen-cells; and in these two cases the stimulating conditions
must be different.

When the eggs have been laid, and the grubs hatched,
the worker bees assume new duties—the feeding and tending
of the young. They eat honey and pollen, which is partially
digested, and supplied as pap to the grubs in such quantities
that they seem bathed in it; but after a short time a mixture
of honey pollen and water is substituted for this pap. It is
said that the drone larve are fed with pap for a longer period
than the workers ; and the queen larva undoubtedly receives
far more of this pap—or, perhaps, of a still richer nutritive
product, sometimes spoken of as royal jelly—and, indeed, is
supplied therewith throughout larval life. It is generally
believed that this high feeding is the cause of queen-develop-
ment, and that should the queen larve die ordinary worker
larve are fed up, and produce queens nowise dissimilar to those
developed in the royal cells. It is clear, if this be so, that the
behaviour of the nurses decides the difference between the
future queens and working bees—that is to say, the fertile
and the sterile females. In any case, the feeding of the young
by members of the same community is a fact to be specially
noted. It is commonly said that the family is the germ from
which the social community springs; and it may be added
that food-collection or food-administration in some form makes
the difference between the family that coheres and the family
that scatters.

When the larve have been fed, each after its kind, the
workers seal up the cells with lids of pollen and wax ; the larvee
spin cocoons, pass into the pupa stage, and then change to perfect
bees, which bite a way through the lid and take their place
in the hive. These young bees now become the nurses, while
the older bees go abroad to fetch honey and pollen to be stored
away in some of the cells. But when a queen emerges, her

SOCIAL COMMUNITIES OF BEES AND ANTS. 209

first act is to go round to the other royal cells, tear them open,
and sting to death the helpless occupants. Meanwhile the old
queen may have led off the surplus population in a swarm, and
the new queen reigns in her stead. Idle drones have also been
emerging from their cells; and when the young queen starts
forth on her nuptial flight she is followed by the drones, mates
with one of them, and returns a potential mother of thousands.
So long as there is abundance of food the useless drones are
tolerated ; but when there is scarcity they are ejected, and
drone eggs, larvee, and pupee are said to be destroyed.

In the works of Huber and others, further marvels of hive-
life, some well-authenticated, others more or less doubtful, are
duly set forth. But enough has here been said to show that
a social community of bees presents problems of animal be-
haviour which are sufficiently difficult of explanation. How
far is the behaviour instinctive? How far is it due to ex-
perience individually acquired ? Are we constrained to admit
a rational factor? If so, is it, like human reason, the result
of generalization from experience of the relationships of
phenomena? Or are there features of insect psychology which
differ from any of which we have firsthand knowledge? These
questions are more easily put than answered. As in the case
of bird-migration, so too in that of the social life of bees,
there is much that honesty forces us to confess our inability
satisfactorily to explain. ;

So, too, is it in the social life of ants. Among these insects
the males and perfect females bear wings, though these appen-
dages may be subsequently shed. In some kinds, however, there
are also wingless males or females capable of exercising the re-
productive function. The workers are wingless, and are often of
two or three kinds, differing in form and appearance, and in some
cases playing different parts in the social economy. There is also,
in some cases, a separate class of large-headed soldier ants ; so
that differentiation of structure among the sterile females is
carried further in ants than in bees. Their nests generally
consist of an elaborate system of chambers and passages, either

built with pine-needles, as in our common wood ant, or
P

210 SOCIAL BEHAVIOUR

hollowed out in the earth or in wood, or sometimes built with
a paper-like material, or formed of rolled leaves. It is said
that a common ant in Eastern Asia (Qeophylla smaragdina)
“forms shelters on the leaves of trees, by curling the edges of
leaves and joining them together. . . . The perfect ant has
no material with which to fasten together the edges it curls ;
its larva, however, possesses glands that secrete a supply of
material for it to form a cocoon with, and the ants utilize the
larvee to effect their purpose.” * This has recently been con-
firmed by Mr. E. G. Green, Government entomologist, at the
Botanic Gardens at Peradeniya, Ceylon. “He has seen ants
actually holding larve in their mouths and utilizing them as
spinning machines. To find out what would be done, some leaves
were purposely separated by Mr. Green. The edges of the leaves
were quickly drawn together by the ants, and, about an hour
later, small white grubs were seen being passed backwards and
forwards across the gaps made in the walls of the shelter. A
continuous thread of silk proceeded from the mouth of the
larva, and was used to repair the damage.” + This is a re-
markable act of apparently intelligent behaviour. But when
we remember how much of the time of ants is occupied in
carrying about their larvae, it is hardly an act of which it
can be affirmed that it could not arise as the result of chance
experience.

In some cases two different genera are found in the same
nest, with separate chambers and passages, as in the case of
the robber-ant (S»/enopsis) and the slave-ant (Mormica fusca).
The orifices by which the former enter are too small to allow
of the entrance of the latter, “hence the robber obtains an
easy living at the expense of the larger species,” for “they
make incursions into the nurseries, and carry off the larve
as food.” é

In a few cases the foundation of a new colony has been
carefully watched. Blockmann was successful in observing
the formation of new nests by Componotus ligniperdus at

* Sharp, “ Insects,” part ii., p. 147.
t Nature, vol. 1xii., p. 253 (July 12, 1900).

SOCIAL COMMUNITIES OF BEES AND ANTS arr

Heidelberg. “He found under stones, in the spring, many
examples of females, either solitary or accompanied only by a
few eggs, larve, or pups. Further, he was successful in getting
isolated females to commence nesting in confinement, and
observed that the ant that afterwards becomes the queen, at
first carries out by herself all the duties of the nest. Beginning
by making a small burrow, she lays some eggs, and when these
hatch, feeds and tends the larva and pups: the first speci-
mens of these latter that become perfect insects are workers
of all sizes, and at once undertake the duties of tending the
young and feeding the mother, who, being thus freed from
the duties of nursing and of providing food while she is her-
self tended and fed, becomes a true queen-ant. Thus it seems
established that, in the case of this species, the division of
labour found in the complex community does not at first exist,
but is correlative with increasing numbers of the society.
Further observations as to the growth of one of these nascent
communities, and the times and conditions under which the
various forms of individuals composing a complete society
first appear, would be of considerable interest.” *

The queen does not, as in the case of the bee, deposit her
eggs in separate cells where they are tended by nurses. The
eggs, which are laid in the chambers of the nest, are subjected
to much licking by the nurses ; the larvee are, moreover, moved
-about from place to place, so as to be subjected to the requisite
conditions of moisture and temperature. They are carefully
cleaned, and after they have passed into the pupa stage the
emerging insects are stripped of a delicate investing skin.
And not only do the ants assiduously feed their young ; those
who have gone forth and drunk their fill of sweet juices feed
those who have remained behind. Forel took some specimens
of Componotus ligniperdus, “and shut them up without food
for several days, and thereafter supplied some of them with
honey, stained with Prussian blue; being very hungry, they
fed so greedily on this that in a few hours their hind bodies

* The quotation is from “ The Cambridge Natural History,” vol. vi.,
“ Insects,” part ii, by David Sharp, F.R.S.; see pp. 145, 146.

212 SOCIAL BEHAVIOUR

were distended to three times their previous size. He then
took one of these gorged individuals, and placed it among
those that had not been fed. The replete ant was at once
explored by touches of the other ants and surrounded, and food
was begged from it. It responded to the demands by feeding
a small specimen from its mouth, and when this little one had
received a good supply, it in turn communicated some thereof
to other specimens; while the original well-fed one also
supplied others, and thus the food was speedily distributed.
This habit of receiving and giving food is of the greatest
importance in the life-history of ants.” * It affords the basis
or starting-point of the keeping of aphides, the making of
slaves, the curious development of honey-pot ants, and in some
cases the association with ants of other insects.

Some of these insects, of which there are many species
belonging to several orders, are parasitic ; others appear to be
hostile, and yet are able to maintain themselves in the nest ;
others simply live side by side with the ants, which seem to be
neither hostile nor friendly to them. In some of these cases
the biological purpose of the association is unknown, while in
others the ant serves as a model which the associated insect
mimics. Thus in the nest of an Indian ant (Sima rufa-nigra)
occur a small wasp and a spider which, to some extent in form
and more markedly in coloration, mimic their hosts. ‘‘ Where-
ever you find this species in any numbers,” says Mr. Rothney,t
“if you watch a few moments, you will see a mimicking spider,
Salticus, running about among the ants, which it very closely
resembles in appearance, much more so in life than in set
specimens placed side by side ; I have seen numbers on the
most friendly footing with the ants, though I have never seen
them enter their burrows. ... They are, I should say, the
only friends the ant has, with the exception of a sand-wasp, a
new species of Ahinopsis since described by Mr. Cameron,
which also very closely mimics the ant, and which, on first

* Sharp, op. ett., p. 147.
+ G. A. J. Rothney, “Notes on Indian Ants,” Trans. Ent, Soe., 1889,
_ p. 354,

SOCIAL COMMUNITIES OF BEES AND ANTS 213

observing among the workers, I took to be the male.” But
there are some beetles which are not only tolerated, but fed
by the ants with which they live. In the case of the genera
Atemeles and Lomechusa, which are always found in or near
ants’ nests, the good offices are reciprocal, for the beetles
“have patches of yellow hairs, and these secrete some substance
with a flavour agreeable .to the ants, which lick the beetles
from time to time. On the other hand, the ants feed the
beetles ; this they do by regurgitating food, at the request of
the beetle, on to their lower lip, from which it is then taken
by the beetle. The beetles in many of their movements
exactly resemble the ants, and their mode of requesting food,
by stroking the ants in certain ways, is quite ant-like. So —

Fic. 25.—Beetle soliciting food from Ant (after Wasmann).

reciprocal is the friendship, that if an ant is in want of food
the beetle will in its turn disgorge for the benefit of its host.
The young of the beetles are reared in the nests by the ants,
who attend to them as carefully as they do to their own young.
The beetles are, however, fond of the ants’ larvee as food, and,
indeed, eat them to a very large extent, even when their own
young are receiving food from the ants. Wasmann (to whom
‘ve are indebted for most of our knowledge on this subject)
seems to be of opinion that the ants scarcely distinguish
between the beetle larvee and their own young ; one unfortu-
nate result for the beetle follows from this, viz. that in the
pupal state the treatment that is suitable for the ant larve

214 SOCIAL BEHAVIOUR

does not agree with the beetle larve. The ants are in the
habit of digging up their own kind, and lifting them out and
cleaning them during their metamorphosis: they do this also
with the beetle larvae, with fatal results; so that only those
that have the good fortune to be forgotten by the ants com-
plete their development.” *

Aphides, or plant-lice, yield to the solicitations of ants,
which stroke them with their antenne, by emitting a drop of
sweet and viscid secretion, and it appears that the caress of the
ant is the natural stimulus for the emission of the drop. Not
only, however, do the ants go forth in search of aphides in
their natural haunts, they bring them to the neighbourhood of
the nest, and may even impound them by building a wall of
earth round and over them. Huber stated that ants collected
the eggs of the aphides and tended them in their nests, and
the accuracy of the observation has been shown by Jord
Avebury and others. “The aphid eggs are laid early in
October, on the food plant of the insect. They are of no direct
use to the ants, yet they are not left where they are laid,
where they would be exposed to the severity of the weather
and to innumerable dangers, but broughi into the nests by the
ants, and tended by them with the utmost care through the
long winter months until the following March, when the young
ones are brought out and again placed on the young shoots of
the daisy.” + Dr. McCook noticed that ants, returning from
the trees on which aphides abounded, fed others near the nests,
and he regarded this as a case of division of labour, the
foragers obtaining food for the nurses which remained in or
near the nest.

A further division of labour, carried to lengths which seem
almost absurd, is found in the honey-pot ant of the United
States and Mexico. The juice on which these ants feed is
obtained from an oak-gall. Foragers go forth at night and
return distended with the sweet fluid, and, having fed the

* Sharp, op. cit., p. 226.
t+ Lord Avebury (Sir John Lubbock), quoted in Romanes’ “ Animal
Intelligence,” pp. 62, 63.

SOCIAL COMMUNITIES OF BEES AND ANTS 215

ordinary workers in the nest, apparently discharge the balance
of their store into living honey-pots, which remain in the nest
and preserve the food till it
may be required by the mem-
bers of the community. Their
abdomens are enormously dis-
tended, they never leave the
nest, and they seem to form
a distinct caste, whose function
it is to passively accumulate
stores of reserve food for the Fig. 26.—Honey-pot Ant.
community. Curiously enough

the same peculiar social arrangement is found in different
genera living as far apart as Mexico, Australia, and South
Africa.

There is no doubt that in some cases the division of labour
is not restricted to the individuals of the same species, but that
other species are introduced into the nest to perform certain
functions—thus giving rise to the so-called slavery among
ants. This is carried to an extreme in the European species
Formica rufescens, the males and queens of which do no work,
while the sole function of the workers is to capture slaves of
the smaller species Formica fusca. In association with this
specialized mode of instinctive behaviour, “even their bodily
structure has undergone a change; their mandibles have lost
their teeth, and have become mere nippers, deadly weapons
indeed, but useless except in war. They have lost the greater
part of their instincts : their art—that is, the power of building ;
their domestic habits—for they take no care of their own young,
all this being done by the slaves ; their industry—they take no
part in providing the daily supplies ; if the colony changes the
situation of its nest, the masters are all carried by the slaves to
the new one ; nay, they have even lost the habit of feeding. . . .
I have had a nest of this species under observation for a
long time, but never saw one of the masters feeding. I have
kept isolated specimens for weeks, by giving them a slave for
an hour or two a day to clean and feed them, and under these

216 SOCIAL BEHAVIOUR

circumstances they remained in perfect health, while, but for
the slaves, they would have perished in two or three days.” *

In this matter, we have in different species successive stages
in the development of the instinctive behaviour which is thus
carried so far in Formica rufescens. Our English ants, of the
species Formica sanguinea, have fewer slaves and are less
dependent on them ; they can feed and forage for themselves,
and during migration carry their slaves—which are of the
same species as in the other case—instead of being carried by
them. In the nests of the common wood ant or horse ant
(Formica rufa) there are occasionally a few slaves. Lord Ave-
bury thinks it likely that they are developed from larve or
pup, originally taken for food, which have by chance come to
maturity in the nest of their captors.

But one more incident in the social life of ants can here be
noticed—though many others could be given did space permit.
The leaf-cutting ants of America form paths from their nests
to suitable trees, from which to obtain the small coin-like leaf
fragments, which they carry in the mandibles, and hence have
gained the name of umbrella or parasol ants. These paths are
sometimes underground ; and Mr. McCook measured one which
ran at a depth of some 18 inches beneath the surface for
448 feet, and was then continued for another 185 feet to’ the
tree which the ants were stripping. The whole path was in an
almost perfect straight line from nest to tree. The leaf
fragments are stored in large quantities in the nest, and it was
long a matter of uncertainty for what purpose they were
collected. The problem was solved by Alfred Mdller, who
found that the leaves, which are subdivided and masticated by
a special set of workers within the nest, form the appropriate
material in which the threads of a fungus ramify and flourish.
This fungus is tended by the ants with great care, and is made
to produce a specially modified form of growth, not found
under other circumstances, in the form of white ageregations,
termed by Moller “Kohlrabi clumps.” These form the
principal food of the ants; and the spongy mass of earth and

* Lord Avebury (Sir John Lubbock), “ Scientific Lectures,” pp. 78, 79.

SOCIAL COMMUNITIES OF BEES AND ANTS 217

leaves is called the fungus garden. ‘If a nest be broken into
and the fungus garden scattered, the ants collect it as quickly as
possible, especially the younger parts, taking as much trouble
over it as over the larvae. They also cover it up again as soon
as possible to protect it from the light.” *

Again, it may be asked with regard to the social life of ants
as with respect to that of bees—How far is their complex
behaviour instinctive ? How far is it due toimitation ? What
part does intelligence play, and under what conditions of
acquisition ? Is reason, in the restricted sense of the word, a
factor in the development of the behaviour ? I cannot answer
these questions, and am of opinion that much detailed observa-
tion is yet needed before we can do much more than speculate
in the matter. Much indeed has been done, but yet more
remains for future investigation.

The conditions under which much of the behaviour is
carried out seem to indicate strong instinctive tendencies
which give an hereditary trend to the direction which the
social behaviour takes. Dr. Bethe,t indeed, goes so far as to
regard the behaviour as almost entirely instinctive, affording
little evidence of that modifiability of reactions which indi-
cates intelligent guidance. He shows as the result of careful
experiment that the behaviour of ants to friends and enemies
are direct reactions to smell. Enemies washed with the excre-
tions of members of the nest are treated as friends, notwith-
standing their different colour, size, and general appearance.
By scent, too, they follow the lead of others and retrace their
way to the nest ; this, he says, is not the result of a mental
process, but is the reaction of a complicated reflex mechanism.
As the outcome of careful observation, Dr. Bethe’s conclusions
are of great value and interest. But he seems to go too far
in denying to ants any power of intelligent accommodation to
circumstances. Jf we admit intelligence, then the fact that

* Nature, vol. xlvii., p. 393 (Aug., 1893), where A. Modlle1’s
investigations are described by J. C. Willis.

+ A. Bethe, “ Diirfen wir den Ameisen und Bienen psychische Quali-
taten Zuschreiben,” Pfliiger’s Archiv., 1xx., 1898.

218 SOCIAL BEHAVIOUR

the insects come forth in the midst of a community in full
social activity would tend to the imitative or intelligent acqui-
sition of like modes of procedure. It is difficult to distinguish
the share taken by these two factors which may well co-
operate. And if natural selection is exercising its influence
through the elimination of those which do not fall into line
in social behaviour, there would be ample opportunity for the
survival of coincident variations.* If one may be allowed to
speculate, it seems probable that the interaction of instinct
and intelligence will be found with fuller knowledge to suffice
for the explanation of the facts, without calling in the known
but here improbable factor of rationality or any factors un-
known elsewhere in psychology.

Some interesting observations of Lord Avebury’s are some-
times quoted as evidence that ants are lacking in intelligence,
but (if we accept the distinction already drawn t) they seem
rather to show the lack of reason. “I placed food,” he says, ¢
“in a porcelain cup, on a slip of glass surrounded. by water,
but accessible to the ants by a bridge, consisting of a strip of
paper two-thirds of an inch long and one-third wide. Having
then put an ant (Formica nigra) from one of my nests to this
food, she began carrying it off, and by degrees a number of
friends came to help her. When about twenty-five ants were
so engaged, I moved the little paper bridge slightly, so as to
leave a chasm just so wide that the ants could not reach
across. They came to the edge and tried hard to get over,
but it did not occur to them to push the paper bridge, though
the distance was only about one-third of an inch, and they
might easily have done so. After trying for about a quarter
of an hour they gave up the attempt and returned home.
This I repeated several times. Then, thinking that paper
was a substance to which they were not accustomed, I tried
the same with a bit of straw one inch long and one-eighth of
an inch wide. The result was the same. Again, I placed
particles of food close to and directly over the nest, but

* See p. 37. t Supra, p. 138.
t ‘‘ Scientific Lectures,” pp. 80-82.

SOCIAL COMMUNITIES OF BEES AND ANTS 219

connected with it only by a passage several feet in length.
Under these circumstances it would be obviously a saving of
time and labour to drop the food on to the nest, or at any rate
to spring down with it, so as to save one journey. But though
I have frequently tried the experiment, my ants never adopted
either of these courses. I arranged matters so that the glass
on which the food was placed was only raised one-third of an
inch above the nest. The ants tried to reach down, and the
distance was so small that occasionally, if another ant passed
underneath just as one was reaching down, the upper one
could step on to its back, and so descend; but this only
happened accidentally, and they did not think of throwing the
particles down, nor, which surprised me very much, would
they jump down themselves. I then placed a heap of mould
close to the glass, but just so far that they could not reach
across. It would have been quite easy for any ant, by moving
a particle of earth for a quarter of an inch, to have made a
bridge by which the food might have been reached, but this
simple expedient did not occur to them.”

Now, when we remember that the method of intelligence is
to profit by chance experience, while the method of reason is,
with foresight and intention, to adapt means to ends, we shall
see that to move a straw even a quarter of an inch, or to make
a bridge with particles of mould, would require rational and
not merely intelligent powers. Chance experience would not
supply the necessary data to be utilized by intelligence when
repetition had established an association in the conscious
situation. Granting that the ants were intelligent but not
rational, they could not be expected to overcome the difficulties,
simple as they seem to us, which Lord Avebury placed in their
path. Had they been overcome the fact would be more difficult
to explain than the use of a stone tool by the sand wasp, since‘
this could more readily be hit upon by chance experience.
And what these valuable experiments, of which kind more
are needed, seem to show is, that the ant, probably the most
intelligent of all insects, has no claim to be regarded as a
rational being.

220 SOCIAL BEHAVIOUR

IV.—AnIMAL TRADITION

In that interaction between instinct and intelligence which,
when further detailed work has sifted and purified our know-
ledge of the psychology of animal communities, may prove
sufficient to account for the well-established facts, animal
tradition will probably have to be recognized as of no little
importance. When a newly emerging ant or bee, or a young
bird or mammal is born into a community where certain modes
of behaviour are already in full swing, an imitative tendency
of the follow-my-leader type may lead it to fall in line with
the traditional habits. It is said that young ants follow the
older workers about the nest, and are ‘“ trained to a knowledge
of domestic duties, especially in the case.of larve.”” On the
other hand, we have seen that, in certain observed cases, the
queen ant is the solitary starting-point of a new community,
and that the division of labour follows with the increasing
numbers of the newly formed social group; so that, in such
cases, whatever part tradition may play in the later phases of
social life, it cannot afford a sufficient account of the division
of labour in the earlier history of the community. We need,
however, fuller information concerning the continued life-
history of such communities under natural conditions, and as
to how far they remain self-contained without any incorpora-
tion of older members from adjoining nests. In the case of
bees, where the old queen departs with a swarm, there may be
greater continuity of tradition. But how far this is a necessary
factor in social development is at present a matter of con-
jecture. In the herd of mammals and the flock of birds, and
in all the family and social life in these classes of animals, the
example of elders, without any imitation of the higher reflective
type, can scarcely be without its influence on the behaviour of
the young which, one would suppose, would tend to fall in with
the ways which had become traditional in the species. Pro-
fessor Wesley Mills tells us that a mongrel pup, whose psychical
development he carefully watched, showed “ extraordinarily

ANIMAL TRADITION 221

rapid” progress when he was introduced to the society of
other dogs, and was thus subjected to the influence of canine
tradition.

How far this influence extends in animal communities—
how far it is either a necessary or even an important con-
‘tributory factor in the development of certain modes of
behaviour—is at present in large degree a matter of specula-
tion. And the only justification for speculation in science is
that it may open our eyes to modes of influence the range and
limits of whose effects may be submitted to the touchstone of
careful observation, and, if possible, experiment. In this
instance it is rather the indefiniteness of the evidence before
us than its absence that stands in the way of any profitable
discussion of the problem from the evidential point of view.
And this indefiniteness is partly due to the fact that the need
of observation is not realized, because this factor in animal
behaviour has not been distinguished with sufficient clearness.
It is worth while, therefore, to devote a short space to a con-
sideration of the relation of this tradition to instinct and
intelligence with a view to the focussing of observation on the
facts by which it may be further elucidated.

In the first place, it is probable that, as in other modes of
animal behaviour, traditional procedure is founded on an
instinctive basis. This must be an imitative tendency of the
broad follow-my-leader type indicated in the first section of
this chapter. And this would afford wide instinctive founda-
tions, which would owe their hereditary character to the fact
that, under natural selection, those individuals in the com-
munity would survive which fell into line with the adaptive
behaviour of their companions, while those which failed in
this respect would be eliminated as more or less isolated out-
siders, standing apart from the social life. In illustration we
may take a hypothetical case, founded, however, upon observa-
tion. The Rev. 8. J. Whitmee, a missionary in Samoa, believes
that the tooth-billed pigeon of these islands (Didunculus
strigirostris) “has probably been frightened when roosting, or
during incubation, by attacks of cats, and has sought safety

222 SOCIAL BEHAVIOUR

in the trees. Learning, from frequent repetition of the fright,
that the ground is a dangerous place, it has acquired the habit
of building, roosting, and feeding on the high trees; and this
habit is now operating for the preservation of this interesting
bird, which a few years ago was almost extinct.” * Now, in
this case, the young birds which followed the lead of those
who, under experience, had acquired the habit, would stand a
better chance of survival than those who, failing to do so, were
caught napping on the ground. In further illustration, we
may take the case of two species of rats found by Mr. C. M.
Woodford on one of the-‘Solomon Islands. These two species
are regarded by Mr. Oldfield Thomas as slightly altered
descendants of one parent species, with adaptations due to
the fact that, of this original species, some have adopted a
terrestrial, others an arboreal life. Thus MM/us rex lives in
trees, has broad footpads, and a long rasp-like, probably semi-
prehensile tail ; while J/us imperator lives on the ground, has
smaller pads, and a short smooth tail. How far the different
modes of behaviour in the two species may have been fostered
by the influence of tradition we do not know; but it is not
improbable that such an influence would be a co-operating
factor in the process of segregation, and that in the course of
time each form has been adapted to its special environment
through the elimination of those individuals which were not
in harmony with the conditions of their life.

Such a case—admittedly hypothetical in the interpretation
put upon the facts—may help us to see how the general
instinctive follow-my-leader tendency might become specialized
in certain essential lines of racial behaviour, and how, under
natural selection, coincident variations in the line of traditional
acts might become more and more definitely inherited as, at
first, strong instinctive tendencies, and eventually more stereo-
typed modes of instinctive behaviour. This, indeed, may
have been the mode of origin of some of the social instincts.

Reverting, however, to the stage where the general instinc-
tive follow-my-leader tendency is only partly or incompletely

* Proceedings of the Zoological Society, 1874, p. 184.

AMIMAL TRADITION 223

specialized along particular lines of behaviour, we should
have at this stage certain hereditary trends of action, depen-
dent on stimuli afforded by the behaviour of others, but
needing, for their guidance to finer issues and more adequate
and highly perfected performance, the play of intelligence and
the satisfaction of nascent social impulses. In the economy of
the hive or the nest there are, no doubt, instinctive tendencies
and predispositions ; but there is also something more than
organic heredity with its transmitted modes of behaviour
analogous to the inherited form and structure of the body or
its parts. Consciousness exerts a guiding influence. The
insect is not independent of experience, but is capable of
profiting by the teachings of that fertile nother of all intelli-
gent behaviour. It is unnecessary, however, to insist on the
fact that such insects are something more than instinctive
automata, but are guided in their behaviour by the results of
experience. Many careful observers lay stress upon this; if,
indeed, they do not go further and claim for the social insect
the higher rational faculty. “When we see,” says Lord
Avebury,* “an ant-hill tenanted by thousands of industrious
inhabitants, excavating chambers, forming tunnels, making
roads, guarding their home, gathering food, feeding the young,
tending their domestic animals—each one fulfilling its duties
industriously, and without confusion—it is difficult altogether
to deny to them the gift of reason; and the preceding
observations tend to confirm the opinion that their mental
powers differ from those of man, not so much in kind as in
degree.”

If the term “ reason” be here accepted in the broad sense,
and not in the narrower sense before indicated, this passage
will probably be endorsed by the majority of those who have
paid any attention to the subject ; save that those who regard
“reason,” in the more restricted acceptation of the term, as
outside any scheme of evolution, may claim that this does con-
stitute a difference in kind and not merely in degree. In
any case the passage expresses the conviction of a close and

* “Scientific Lectures,” 2nd edit., p. 140.

224 SOCIAL BEHAVIOUR

singularly unprejudiced observer, that the doings of ants in-
volve conscious guidance in the light of experience individually
acquired.

And yet the behaviour of different species of ants, each
after its kind, is remarkably constant—so constant that, to
use the words of Dr. Peckham in another connection, it is
characteristic of the species, and would be an important part
of any definition of the insect based upon its habits. And
some part of this constancy may be due to tradition, though
much of it may result from strong instinctive tendencies which
intelligence guides to similar ends, because the conditions are
similar in successive generations of social insects.

From the point of view of observation, however, it is par-
ticularly difficult to distinguish the part played by tradition as
a psychological influence from that played by what we have
above described as instinctive imitation. In our study of other
modes of instinctive behaviour we can isolate an individual, or
group of young individuals, and observe how far certain acts
are performed prior to any experience. Thus chicks behave in
certain instinctive ways under conditions which preclude their
learning from the hen or other older birds—so that tradition
cannot be operative. But where social behaviour is concerned,
such methods of observation are necessarily excluded—save in
such cases as that of the incipient community of ants. For if
certain instinctive acts require for their due performance the
stimulus of the like performance in others, what is this but a
form of instinctive tradition ; and how are we to distinguish it
from intelligent tradition, where a psychological factor has
freer play and exercises guidance over the performance? In
the present state of our knowledge we can do no more than
suggest, as not improbable, that tradition passes through three
phases: the first in which it is instinctive; the second in
which it becomes intelligent through the satisfaction which the
due performance of traditional acts arouses in consciousness ;
and the third in which, at any rate in man, it takes on a
rational form, and is made to accord with an ideal scheme, the
product of conceptual thought and of reflection on data which

THE EVOLUTION OF SOCIAL BEHAVIOUR 225

have been generalized and considered in their due relationships
to the scheme which takes definite form in the mind. Whether
in the social communities of insects or those of beavers, among
mammals, or rooks among birds, tradition has begun to pass
into the third or rational stage, we do not know. It may be
so, but probably the development along these lines has not
been carried far. Presumably in the ant, rook, and beaver
anything like an ideal scheme of thought based on reflection, if
it exist, is as yet exceedingly indefinite.

But even supposing that no animal has yet risen beyond
the second or intelligent stage, it is none the less important
to realize that we have here, in animal life, the foundations on
which may be raised what may, perhaps, be regarded as one
of the characteristic features of human progress. This charac-
teristic is the transference of evolution from the organism to
the environment handed on from generation to generation.
Thus man, “availing himself of tradition, is able to seize upon
the acquirements of his ancestors at the point where they left
them.” * Thus “he has slowly accumulated and organized
the experience which is almost wholly lost with the cessation
of individual life in other animals.” + But he is able to do
so through the extension, refining, and fixing of that instinctive
and intelligent tradition which begins to take form in animal
communities.

V.—Tue Evonvrtion or Soctan BEHAVIOUR

“Animals of many kinds,” said Darwin,f “are social ;
every one must have noticed how miserable horses, dogs,
sheep, etc., are when separated from their companions. The
most common mutual service in the higher animals is to warn
one another of danger. Every sportsman knows how difficult

* Weismann, “ Essays,” vol. ii, p. 50.

+ Husley, “ Collected Essays,” vol. vii., p. 155.

+ “The Descent of Man,” vol. i. p. 853, 2nd Ed., 1888, The quotations
from Darwin in this paragraph and that which follows are somewhat con+
densed by a few omissions.

Q

226 SOCIAL BEHAVIOUR

it is to approach animals in a herd or troop. Wild horses and
cattle do not, I believe, make any danger-signal; but the
attitude of any one of them who first discovers an enemy
warns the others. Rabbits stamp loudly on the ground with
their hind feet as a signal; sheep and chamois do the same
with their fore feet, uttering likewise a whistle. Many birds
and some mammals post sentinels. The leader of a troop of
monkeys acts as such, and utters cries expressive both of
danger and of safety. Social animals perform many little
services for each other: horses nibble, and cows lick each
other; monkeys search each other for external parasites, and
are said to remove thorns and burrs. Social animals mutually
defend each other. Bull bisons in North America, when there
is danger drives the cows and calves into the middle of the
herd, whilst they defend the outside. Among baboons the old
males come forward to the attack. Wolves hunt in packs;
and pelicans fish in concert.

“Tt has often been assumed,” continues Darwin, “ that
animals were in the first place rendered social, and that they
feel as a consequence uncomfortable when separated from each
other, and comfortable whilst together ; but it is a more pro-
bable view that these sensations were first developed, in order
that those animals which would profit by living in society
should be induced to live together, in the same manner as the
sense of hunger and the pleasure of eating were, no doubt, first
acquired in order to induce animals to eat. The feeling of
pleasure from society is probably an extension of the parental
and filial affections, since the social instinct seems to be
developed by the young remaining long with their parents ;
and this extension may be attributed in part to habit, but
chiefly to natural selection. With those animals which were
benetited by living in close association, the individuals which
took the greatest pleasure in society would best escape various
dangers ; while those which cared least for their comrades, and
lived solitary, would perish in greater numbers. In however
complex a manner the feeling of sympathy may have originated,
as it is one of high importance to all those animals which aid

THE EVOLUTION OF SOCIAL BEHAVIOUR 227

and defend one another, it will have been increased through
natural selection ; for those communities which included the
greatest number of the most sympathetic members would
flourish best, and rear the greatest number of offspring.”

It is impossible to improve upon this pithy description
of the salient facts, and terse explanation in terms of the
hypothesis of natural selection. It may, perhaps, be urged
that, on this hypothesis, the origin of the social state, through
a biological association of individuals, probably neither pre-
ceded nor followed the development of a psychical bond arising
from the sense of satisfaction and comfort afforded by social
life, but that both originated pari passu. If the linkage was
primarily instinctive, its intelligent continuance could only be
effected through the pleasure social behaviour carried with it,
and the discomfort of separation from the community. No
instinctive acts would be persistently repeated, under the
guidance of individual experience, if that experience proved
bitter and not sweet. An animal with thwarted instincts is
one with unsatisfied impulses ; its biological and its psycho-
logical tendencies are alike unfulfilled. What Darwin saw
and wished to enforce, however, was that the psychical link of
conscious satisfaction was a necessary prerequisite of the con-
tinuance and further evolution of sociability ; and that without
the integrating bonds of sympathy any advance of social
development was impossible.

In two able and interesting articles in the Mineteenth
Century review,* on “Mutual Aid among Animals,” Prince
Kropotkine gives a useful and sufficiently detailed summary of
the chief facts concerning the social relationships which have
been observed in the animal kingdom—including, perhaps,
some rather apocryphal instances,—and combats Huxley’s state-
ment + that, “ beyond the limited and temporary relations of
the family, the Hobbesian war of each against all is the normal
state of existence” among animals and primitive men. “ Life

* Vol. xxviii, Sept. and Nov., 1890, pp. 337-354, 699-719.

+ Nineteenth Century, Feb., 1888, p. 165. “ Collected Essays,” vol. ix.,
p. 204.

228 SOCIAL BEHAVIOUR

in societies,” says Prince Kropotkine, “is no exception in the
animal world. It is the rule, the law of nature, and it reaches
its fullest development with the higher vertebrates. Those
species which live solitary, or in small families only, are re-
latively few, and their numbers are limited.”* “Life in
societies enables the feeblest insects, the feeblest birds, and
the feeblest mammals to resist, or to protect themselves from,
the most terrible birds and beasts of prey ; it permits longevity ;
it enables the species to rear its progeny with the least waste
of energy, and to maintain its numbers, albeit with a very
slow birth-rate ; it enables the gregarious animals to migrate
in search of new abodes. Therefore, while fully admitting
that force, swiftness, protective colours, cunningness, and
endurance to hunger and cold, which are mentioned by Darwin
and Wallace, are so many qualities making the individual or
the species the fittest under certain circumstances, we maintain
that under any circumstances sociability is the greatest
advantage in the struggle for life... . The fittest are thus
the most sociable animals, and sociability appears as the chief
factor in evolution, both directly, by securing the well-being
of the species while diminishing the waste of energy, and
indirectly by favouring the growth of intelligence.” t And
summarizing his argument, Prince Kropotkine says,t ‘ We
have seen how few are the animal species which live an isolated
life, and how numberless are those which live in societies,
either for mutual defence, or for hunting and storing up food,
or for rearing their offspring, or simply for enjoying life in
common. We have also seen that, though a good deal of
warfare goes on between different species, or even different
tribes of the same species, peace and mutual support are
the rule within the tribe, or the species; and that those
species which best know how to combine, and to avoid com-
petition, have the best chances of survival and of further

* Op. cit., pp. 709, 710.

+ Page 711.

t “Mutual Aid among Savages.” Ménetecenth Century, vol. xxix.,
April, 1891, p. 538.

’

THE EVOLUTION OF SOCIAL BEHAVIOUR 2:29

progressive development. They prosper, while unsociable
species decay.”

Prince Kropotkine seems, however, to push his argument
too far. The assertion that the fittest are the most sociable
animals, that sociability appears as the chief factor in evolution,
and that unsociable species decay, is not likely to be accepted
without qualification by zoologists. What grounds have we
for saying that the solitary wasps are less fit than the social
wasps? Hach has a fitness according to its kind. Can it be
maintained that the unsocial tiger is less fit than the social
jackal ? And can it be said that tigers, which are reported
absolutely to swarm in Java and Sumatra, exemplify the
decay of an unsociable species? Is it seriously contended that
the hawk, which may be successfully mobbed by a number of
wagtails, is less fit than his more social assailants ? And are
the unsocial raptorial birds decaying species? Such questions
might be asked by the score. And the answer in every case
is that the social and unsocial alike are fitted to their several
states of life. In fact, it might be contended, with every
whit as much if not more cogency, that sociability is nature’s
device for enabling the weaker, and hence in themselves the
less fit, to resist the attacks and encroachments of the stronger
and individually fitter. Discussing the possibilities of human
ancestry, Darwin said : * “In regard to bodily size or strength,
we do not know whether man is descended from some com-
paratively small species like the chimpanzee, or from one as
powerful as the gorilla. We should, however, bear in mind
that an animal possessing great size, strength, and ferocity,
which, like the gorilla, could defend itself from all enemies,
would not perhaps have become social ; and this would most
effectually have checked the acquirement of the higher mental
qualities, such as sympathy and the love of his fellows. Hence
it might have been an immense advantage to man to have
sprung from some comparatively weak creature.”

Zoologists, again, will hardly accept without question
Prince Kropotkine’s assertion that “life in societies is no

* “ Descent of Man,” vol. i., p. 96.

230 SOCIAL BEHAVIOUR

exception in the animal world, but is the rule, the law of
nature.” Many will contend, on the other hand, that life in
societies with anything like division of labour, or with mutual
aid (and this seldom carried far), is, taking the animal kingdom
as a whole, of comparatively rare occurrence, though none the
less noteworthy where it exists. Aud, in any case, it seems
somewhat extravagant to say that sociability is the chief factor
in evolution. No doubt it might be plausibly urged that
human society is, from man’s point of view, the highest
product of evolution ; that in attaining to this end sociability
has been the leading factor ; and that obviously the leading
factor in the evolution of the highest product may properly
be called the chief factor in evolution. But Prince Kropotkine
apparently means that sociability is the chief factor, not only
in this evolution, but in all organic, or, at least, all animal
evolution. In this he will receive the support of but few
zoologists. By some extravagance of statement he has
weakened his own case, which is otherwise not lacking in
points of weakness. The legitimate inferences from animal
behaviour are, that co-operation is in some cases a factor in
the evolution of a successful species, that in human progress
it has been an important factor giving strength to a creature
weak in tooth and claw, and that this factor has co-existed,
and still coexists, with that of competition, in the absence of
which the race would be dragged down to lower levels of
efficiency by the incubus of weaklings.

To Professor Alfred Espinas * we owe the best and fullest
discussion of the social life of animals, and to his work the
reader may be referred for a careful and, for the most part,
unstrained and unbiassed consideration of the phenomena.
In common with others who have devoted serious attention
to the subject, he sees in the family the starting-point of the
higher and more comprehensive social group, or “ peuplade.”
Prince Kropotkine seems, indeed, to combat this view ; but
the divergence of opinion is more apparent than real. He

* “Deg Sociétés Animales: tude de Psychologie Comparée” (Paris,
1877).

THE EVOLUTION OF SOCIAL BEHAVIOUR 231

tells us * that anthropology “has established beyond any doubt
that mankind did not begin its life in the shape of small
isolated families. Far from being a primitive form of
organization, the family is a very late product of human
evolution. ... Societies, bands, or tribes—not families—
were thus the primitive form of organization of mankind
and its earliest ancestors.” And in support of his views he
adduces the sexual communism which is said to be found in
the lowest savages, and briefly traces the development of
monogamy and the genesis of the family ideal as we conceive
it. It may at once be admitted that in all probability man-
kind did not have its origin in small isolated families. If we
do not admit this we must accept the alternative hypothesis,
that man was developed from an unsocial ancestor. For
though the biological family is the starting-point of the
community, it does not of course follow that wherever there
is so much coherence between parents and offspring as to form
a temporary family group, a social community must in due
course arise. In such unsocial carnivora as the tiger, the
temporary linkage of family life is strong while it lasts. But
though mankind presumably originated in a prebuman race
that had already reached some degree of social coherence,
there remains behind the question—what was the origin of
this social group? And to this question, Prince Kropotkine,
in common with Darwin and Espinas, would probably answer
without hesitation, that the primeval germ of the social com-
munity lay in the prolonged coherence of the group of parents
and offspring. In the unsocial animals the family separates
and disintegrates before the offspring mate. But if the family
continue to cohere, the mating of offspring will give rise to the
continuity of coherence found in the herd, or troop, or tribe.
For new family groups will be constantly arising before the
old family groups have ceased to be associated. Thus would
be afforded more opportunity for tradition than among the
unsocial animals.

* “Mutual Aid among Savages,” Nineteenth Century, April, 1891, pp.
539, 040.

232 SOCIAL BEHAVIOUR

e

How, then, can it be said that, “far from being a primitive
form of organization, the family is a very late product of
human evolution”? By using the word “family” in a sense
somewhat different—nay, widely different—from that in which
it is employed in a biological discussion. In the latter usage
sexual communism is not excluded ; A., B., and C., D. may have
offspring this season; A., D., and C., B. next season. In each
season there are family groups with interchange of partners. This
does not, however, conform with our conception of the family
as realized under civilization. Herein, in fact, lies the essential
difference between the human and the animal family. The
one is a realized ideal ; the other is merely a natural occurrence.
Even in the case of monogamous animals, mating for life is
probably not conduct in conformity with an ideal, but is due
to the fact that instinctive tendencies have taken this line of
direction. On the other hand, in monogamous communities of
mankind, there is, unfortunately, evidence that in some cases
the ideal is not strong enough to prevent presumably ancestral
tendencies in the direction of communism.

The basis of human social conduct is unquestionably to be
traced in the social behaviour of animals, in inherited tendencies
to co-operation and mutual help, in the bonds of sympathy
arising through the satisfaction of impulses towards such
behaviour, and perhaps, to some extent, in the influence of
tradition. It is not, however, until this tradition is rendered,
through descriptive communication, more continuous and more
effective ; it is not until an ideal of mutual aid, and social
conduct generally, takes form and is rendered common to the
tribe; it is not until the more or less realized conceptions of
one generation are handed on to become the environment
under which the succeeding generations are nurtured ; it is
not indeed until man consciously and reflectively aims at the
bettering of his environment in accordance with standards
rationally conceived and deliberately carried into execution ;
that a new régime of civilized progress, elsewhere unknown in
nature, takes definite form. Under this régime, the elimination
of failures through natural selection, though it may not be

THE EVOLUTION OF SOCIAL BEHAVIOUR 233

entirely superseded, plays a subordinate part ; alongside the
organic continuity which is due to physical heredity, there runs
a continuity of tradition through social inheritance.

Human civilization is an embodiment of reason, a product
of reflection, a realization of ideals conceived by the leaders of
mankind. All this forms the environment of each one of us.
And it is this environment which is undergoing progressive
evolution and playing on the rational faculties of those which
are submitted to its moulding influence. There is no sufficient
evidence of anything of the kind in the social communities of
animals. This, of course, must be accepted merely as an
expression of opinion. But on the hypothesis that animals
are rational beings, capable of reflection, it is difficult to
understand why they should remain at so low a level of social
achievement. The absence of powers of descriptive intercom-
munication is often assigned as the cause of their comparatively
unprogressive condition ; but it may be regarded as the sign,
rather than the cause, of their lack of reason in the more re-
stricted sense of the term. We cannot, however, enter into
the much-disputed question whether reason is the product of
language, or language the outcome of reason. Perhaps the
safest position is to assume that rationality and true speech are
in large measure different aspects of one evolutionary move-
ment—speech arising out of such preceding modes of communi-
cation as were considered in the second section of this chapter ;
reason developing out of intelligence which supplies its neces-
sary data. It is sometimes said that, notwithstanding their
powers of speech, savages in their social relations show little
advance on animal communities. But surely such statements
must be made in forgetfulness of the fact that savage customs
almost invariably indicate the presence and sway of ideals which
puzzle us from their quaintness, and from the fact that they
seem contrary to the dictates of intelligence and due to motives
and conceptions the nature and force of which we find it diffi-
cult to estimate. The passage from intelligent social behaviour
to the rationality which has assumed such strange aspects among
existing savages took place somewhere at some time in the past ;

234 SOCIAL BEHAVIOUR

and the stages of its evolution are hidden from our view.
All we can say is, that it is possible to trace in animal be-
haviour some of the instinctive tendencies and intelligent modes
of accommodation to social circumstances, together with the
germs of imitation, intercommunication, and tradition, and the
establishment of bonds of sympathy, without which the subse-
quent stages of evolution would be inconceivable.

CHAPTER VI

THE FEELINGS AND EMOTIONS

I—Inevunss, Interest, anp Emorron

Any discussion of animal behaviour must deal largely with
what is termed the conative aspect of consciousness. “The
states designated by such words as craving, longiny, yearning,
endeavour, effort, desire, wish, and will,” says Dr. Stout, in his
admirable “ Manual of Psychology,” * “ have one characteristic
in common. In all of these there is an inherent tendency to
pass beyond themselves and become something different. This
tendency is not only a fact, but an experience ; and the peculiar
mode of being conscious, which constitutes the experience,
is called conation.” Closely associated with this conation is
impulse, which Dr. Stout defines as “any conative tendency,
so far as it operates by its own isolated intensity, apart from
its relation to a general system of motives. Action on impulse
is thus contrasted with action which results from reflection and
deliberation.” | In the interpretation we have advocated,
animals are essentially creatures of impulse, and not to any
large extent, if at all, reflective agents. And their impulses
may be associated either with their inherited and congenital
behaviour, or with that which is due to acquired experience.
Tn other words, their impulses may be divided broadly into
two classes, the one instinctive, the other acquired.

Dr. Stout says that conation is not only a fact, but an ex-
perience. Now, first as to the fact. It seems to be the

* Page 63.
+ Op. cit., p. 267; of. supra, p. 138.
235

236 THE FEELINGS AND EMOTIONS

correlative in consciousness of the behaviour of the nervous
system under stimulation. Let us take some simple case such
as that, for example, of a hungry chick pecking at a grain of
corn. This is explained from the physiological point of view
by saying that internal and external stimulation—internal from
the digestive organs and system in need of food, external
through the eye—gives rise to a state of unstable equilibrium
in the nerve-centres ; and that, when the instability reaches a
certain value, the nervous system discharges into the motor
organs, the chick pecks, and stable equilibrium is restored.
The tendency to discharge in some way under stimulation is
an essential characteristic of a nervous system. It is one of
the facts of physiological science. So, too, the conative
tendency is one of the facts of psychological science ; it is a
change in the situation introduced by the effects of the physio-
logical discharge. :

Let us here parenthetically notice that the physiological
tendency in the nervous system is an evolved complication and
a specialized development of one of the fundamental properties
of protoplasm—that which is often spoken of as irritability.
One of the characteristics of all living matter is its ‘‘ explosive”
instability. So that at the very threshold of organic behaviour
we have the analogue of that which, in its developed form,
becomes the tendency of the nervous system to discharge as
the result of stimulation. The conative tendency of the
psychologist, therefore, has its roots deep down in the elemental
germs of all organic life.

And this conative tendency is, says Dr. Stout, not only a
fact but an experience. Let us return to our hungry chick
that pecks at a grain of corn. And let us grant that as the
result of stimulation there arise states of consciousness which
we describe as a feeling of hunger and the sight of the grain
of corn. The nervous system now discharges ; and there are
introduced into the situation a further group of data, the
motor consciousness of the actual behaviour, sensory data
from the results of the act, the seizing of the grain and so
forth. The situation has unquestionably changed. But ig

IMPULSE, INTEREST, AND EMOTION 237

there any specific consciousness of the conative tendency as
such? Is there any “peculiar mode of being conscious
which constitutes the experience which is called conation ” ?
It is difficult to say. Hence we find differences of opinion
among psychologists as to what, from the psychological
point of view, the impulse actually is. Is it simply the
conscious situation prior to the response? Is it a feeling
of the change from the initial to the succeeding phase? Or
are new data introduced apart from those afforded by stimula-
tion on the one hand and response on the other? We will
not attempt to decide. Without determining its exact nature
we may rest content with the very general statement that
impulse is a concomitant of a change in the conscious situation.

There is, however, a use of the term concerning which it
seems necessarily to enter a word of protest. Impulse is by

- some regarded as the underlying cause of the conative tendency.

Now science, as such, has nothing whatever to do with under-
lying causes. If, as a matter of observation and inference, we
have reason to believe that there is such a tendency, science
simply accepts the fact, and endeavours to formulate the con-
ditions under which it arises, and to trace its observed or
inferred antecedents. No doubt many of us find it difficult or
impossible to rest content with the strictly scientific position,
that of unquestioning acceptance of the facts of nature as we
find them given in experience. We say: Here is an observed
tendency the conditions and antecedents of which are described
by science. But what causes the tendency ; what is the im-
pelling force? Now to such questions science can give no
answer. Science deals with phenomena, and tries to tell us all
about their conditions and their antecedents. But whenever
Science is asked : “ What is the underlying cause of the pheno-
mena,—that which calls them into being?” Science should
always give one answer and one only: ‘Frankly, I do not
know; that lies outside my province; ask my sister Meta-
physics.” Science ought to have nothing whatever to do with
force as the underlying cause of anything in this universe of
phenomena. And impulse, as the impelling force which calls

238 THE FEELINGS AND EMOTIONS

a conative tendency into being, is a metaphysical, not a
scientific conception.

We need not further discuss the psychological nature of
impulse. Indeed, the little that has been said would not have
been necessary to our inquiry were it not that we frequently
have occasion to speak of animals as “ creatures of impulse,”
and to refer to their behaviour as due to impulse. What do we
mean by such expressions ? If we regard conative tendency
as a fact (whatever may be said for or against its being also a
specific experience), and if this fact is the tendency of the
conscious situation to develope in certain definite ways, then
we may define impulse with sufficient clearness by saying,
with Dr. Stout, that it is characterized by being unreflective.
Conative tendency thus comprises two categories—impulse and
volition ; the one unreflective, the other involving deliberation.

Before passing on to consider how impulse is partly
determined by the feeling-tone and the emotional attributes
of the conscious situation, we may first draw attention to the
important way in which the results of conative tendency
afford the data through which consciousness attains its unity
in the midst of diversity of experience.

We said that the impulses might be divided broadly into
two classes—the one instinctive, the other acquired. Now,
from the point of view suggested by a study of behaviour, if
not also, as I am disposed to think, from the more general
standpoint of a genetic study of mental development, it is
convenient to start with the instinctive act and the conscious
situation it implies. We have here a piece of experience
which, if we may so phrase it, hangs together; in which
experience of things in the environment is included in the
same elemental synthesis with that of bodily acts in organic
relation to these things. It is closely linked, on the one hand,
with a foregoing act of attention, itself of the instinctive type ;
closely linked, on the other hand, with the results of behaviour
through which the environing things call forth a new conscious
situation and evoke a further response. Thus not only does
the experience of an instinctive act hang together, but a series

IMPULSE, INTEREST, AND EMOTION 239

of such acts do so likewise. And coalescent association not
only links and groups the elements within the situation called
forth by the single act, but comprises also the elements of the
developing situation afforded by the whole series. We see this
in the young chick, where, as the result of experience, attention
is emphasized where the material is palatable, and lapses where
it is nauseous—such nauseous substances being soon ignored.
Furthermore many environing things appeal in different ways
to the same limited number of sense organs, while the same
motor organs respond in different ways in successive modes
of instinctive behaviour. The same brain forms the physical
basis of varied situations overlapping in many ways, and
receives afferent messages from the same body. Hence, in
its organic unity it affords the conditions for an underlying
stratum of mental unity, amid all the diversities of experience ;
while the multiplicity of messages on the one hand from
external things, and on the other hand from internal happen-
ings, lays the foundations of a differentiation between the
external world and the self—a differentiation long to remain
implicit, and only to be rendered explicit on a far higher level
of mental development. For at this early stage, and perhaps
throughout animal life, “there is no single continuous self con-
trasted with a single continuous world. Self, as a whole,
uniting present, past, and future phases, and the world as a
single coherent system of things and processes, are ideal con-
structions, built up gradually in the course of human develop-
ment. The ideal construction of self and the world is com-
paratively rudimentary in the lower races of mankind, and it
never can be complete. On the purely perceptual plane [with
which we are now dealing] it has not even begun.”* But
though the ideal constructions of self and the world have not,
as Dr. Stout says, at this stage, even begun, yet, as the same
author observes,t “animals distinguish in the environment, and
‘treat as a separate thing, whatever portion of matter appeals to
their peculiar instincts, and affords occasion for their charac-
teristic modes of activity.” And this differentiation of specially
* G, F. Stout, “ Manual of Psychology,” p. 268. t Ibid., p. 318.

240 |. THE FEELINGS AND EMOTIONS

interesting things from each other, and from their relatively
uninteresting surroundings, must be accompanied by some
differentiation of these things from themselves as affected by
them and reacting to them. So that here, as we have seen
to be the case in other matters, what is commonly called the
perceptual life of animals affords the rough-hewn materials
from which ideal constructions may be elaborated by rational
beings.

We cannot here attempt to do more than barely indicate
the manner in which the perceptual process in animals may
acquire unity and diversity—unity through the functioning of
the same brain and body, diversity from the different modes
of functioning and the differential effects of diverse modes
of stimulation. The interesting point for us in our special
inquiry is that it is through behaviour that all this is brought
about.

As we interpret the facts, the restless activity of the young
is primarily a biological fact, and is to be dealt with as an
organic problem—a complication of the fundamental irritability
of protoplasm. But it is also an essential condition to the
acquisition of conscious experience; and the more there is
of it in varied modes the wider is the range of the data
afforded to consciousness. Behaviour is thus the goal of
organic heredity, and the starting-point of conscious accom-
modation and adjustment ; it is the biological end of variation,
and affords the means to intelligent modification.

So much for some of the results of conative tendency.
Not only does it secure adaptation or adjustment to the
environment, but it affords the conditions of mental develop-
ment by which further accommodation is rendered possible.
But, in addition to the attainment of biological ends, in
addition to the furtherance of survival in the struggle for
existence, mental development has another aspect. All sensory
data, whether from the special senses, from the motor processes
concerned in responsive behaviour, or from other sources, may,
and perhaps always do, carry with them some amount of what
is termed feeling-tone, giving rise to a net result in consciousness

IMPULSE, INTEREST, AND EMOTION 241

which we call pleasure or the reverse. Pleasure or satis-
faction—however we name that which, though vague and
indeterminate in outline, is a very real attribute of the con-
scious sitnation—affords its sanction to certain modes of
conation, and may thus be regarded as the psychological end
of their continuance or their repetition. It is partly, no doubt,
a direct adjunct of sight, hearing, taste, and so forth, and of
smooth and easy movements of the body and limbs; but it ig
partly due to a great body of stimulation coming from many
parts of the organic system. The blood-vessels are dilated
or contracted, the heart’s action increased or diminished ;
respiration is deepened or the reverse, and its rhythm may
be altered; glands are thrown into a state of activity ;
the tone of the muscles is affected, and there may be either
incipient contraction or relaxation. These are primarily
organic effects ; but they influence the conscious situation, and
are themselves suffused with feeling-tone. For, from all the
parts so affected, messages are carried in to the brain, and such
afferent messages afford data to consciousness. It may be
that the experience of the conative tendency, for which
Dr. Stout and others claim a distinctive place in consciousness,
is largely due to afferent messages from the motor organs
incipiently innervated in preparation for the behaviour which
follows. In any case these probably form very important
elements in the conscious situation antecedent to the actual
response. In what we may term motor attention—the state
well exemplified by a cat in the strained pause which precedes
the spring on to the prey, or in ourselves when we poise
before a dive or hold the billiard cue in preparation for a
delicate stroke—this incipient innervation, felt through afferent
messages from the parts thus braced for action, enters with
much distinctness into the conscious situation. In sensory
attention, on the other hand, reflex acts have actually taken
place, having for their end and purpose the focussing of the
sense organs on the object which stimulates them, so that in
this way further and more effective stimulation may be received.
But, as the sense organ is steadily held to the focus, and made
R

242 THE FEELINGS AND EMOTIONS

effectually to cover the stimulating thing, the motor apparatus
concerned is kept on the strain, and is all the while con-
tributing data to the conscious situation.

In primary genesis attention, both motor and sensory, is
unquestionably organic and reflex in its nature. It is a pro-
duct, and an invaluable product, of biological evolution.
Without this as a basis, the higher forms of attention under
conscious guidance would be impossible. For all these higher
forms are modifications and complications of what is given in
organic heritage. Here, as elsewhere throughout the whole
range of behaviour, consciousness only guides to finer issues
what is presented to it in rough outline, or in isolated frag-
ments, as the outcome of biological evolution. But the organic
responses afford the data which consciousness uses that it may
mould and fashion the behaviour so as to reach higher and
more complex modes of adjustment.

Lest a familiar form of words should give rise to mis-
apprehension, it may here be stated that, when we say that
consciousness moulds and fashions behaviour, we do not intend
to imply that consciousness is an underlying cause. We are
not using the term consciousness in a metaphysical sense. We
mean that consciousness is the expression of certain conditions
under which behaviour is guided. Instead of saying, there-
fore, that consciousness utilizes certain sensory data, it would
be more correct to say that it is the sum-total of these data
which are the psychical expression of certain brain conditions
under which behaviour, as a matter of fact, takes a given set
or direction. We use the word consciousness, then, not in its
metaphysical sense of an underlying cause or force, but in its
scientific sense, as the concomitant of certain antecedent con-
ditions. Our common modes of speech lend themselves with
misleading facility to metaphysical assumptions, all the more
insidious since they are not consciously acknowledged as such.
And not only what we comprise under the broader group-
name ‘ consciousness,” but what we include under narrower
group-names, such as “impulse,” “volition,” “ instinct,”
“intelligence,” “reason,” and the like, often do duty as

SE

a

IMPULSE, INTEREST, AND EMOTION 243

underlying causes of the phenomena, which, from the scientific
point of view, they do no more than name.

We often say, for example, that interest guides behaviour
in this direction or in that. But such interest must not be
regarded as an impelling force ; it is an attribute of the con-
scious situation, more or less suffused with feeling-tone. It is
not easy to define; but it seems to take on its distinctive
character when re-presentative elements contribute what Dr.
Stout * terms “meaning” to the conscious situation. The
meaning in the early stages of mental development is, however,
merely perceptual, and not that which comes much later—that
which is implied in the phrase “rational significance.” In
the chick which has tasted a cinnabar caterpillar the situation
evoked by the sight of this larva has meaning in virtue of
the actual experience. But, in this case, the meaning is not
conducive to continued interest, since it checks, rather than
stimulates, behaviour. At first, indeed, there may be the
repellent interest of aversion. But this passes by, and the
larvee are soon ignored. Small worms also acquire meaning,
and here the interest is attractive, and is stimulated afresh
each time the meaning is reinforced by repetition of the
act of seizing and swallowing.

We have seen that it is through behaviour that things
become differentiated from their surroundings, and acquire
relative independence in experience. It is through behaviour
that what we have termed conscious situations develop. The
thing is the centre or nucleus of a developing situation—that
which starts the behaviour, and towards which the behaviour is
directed ; or, since the behaviour may be that of avoidance or
escape, we should, perhaps, rather say, it is that to which the
behaviour has reference. Now, if interest is the feeling-tone
attaching to the whole attentive situation, and if the nucleus
of the situation is the thing, it naturally follows that the thing
becomes the centre of interest. The mouse is a centre of
absorbing interest to the cat, her eggs to the mother-bird, his
mate to the sparrow in the spring. Companions are centres

* “ Manual of Psychology,” p. 84.

244 THE FEELINGS AND EMOTIONS

of abiding interest to social animals, because they are also the
centres of social behaviour and the conscious situations arising
thereout ; because they evoke in special ways the attentive
situation.

The differentiated thing being thus a centre of interest, a
relatively fixed nucleus in a changing conscious situation, the
development of which is due to behaviour, there can be no
question that, among social animals, the companion becomes
a peculiar and specialized centre. Around him develops a
particular type of behaviour. Towards him the reactions are
of a quite distinctive kind. Mother and offspring, mate and
mate, are reciprocal centres of interest. To the offspring the
parent is a common centre of interest. As they grow up
together, what is of interest to one is likewise of interest at
the same time to others. Imitation begets similarity of con-
scious situations. In many ways such community of interest
is fostered ; and through this community of interest the con-
scious situations acquire their distinctively social character.
Not only is the companion, as the nucleus of a situation, a
thing which reacts in altogether special ways, so that it becomes
differentiated from other things as something the meaning
of which, and the interest in which, are sui generis and unique
in type; but it enters into other situations in ways that are
also peculiar and characteristic. A worm is thrown to a couple
of chicks, and is to each a centre of interest—the nucleus of
a situation involving appropriate modes of behaviour. But
into this situation there enters for each of them, in a quite
peculiar and distinctive way, the action and behaviour of the
other chick. The situation is complicated by the introduction
of a second centre of interest, and the behaviour has reference
to both centres. Instead of quietly and leisurely dealing
with the worm in accordance with its special meaning, as
it does when there is no rival in the field, the chick darts at it,
and bolts with it in accordance with the special meaning
which its neighbour’s presence, under such circumstances, has
acquired. And this different behaviour carries with it a felt
difference in the conscious situation—the interest of which is

IMPULSE, INTEREST, AND EMOTION — 245

centred in the companion. Or take the case of a herd of
cattle, which attacks a common enemy. The enemy is the
primary nucleus of the situation, but it is profoundly modified
by the presence of companions by which the behaviour of attack
is determined. The situation is social, and not merely indi-
vidual, and a social interest suffuses it, and gives it a distinctive
character.

Tn this social interest probably arise the germs—but only the
germs—of the sense of personality. Some, indeed, go so far as
to urge that we learn to know ourselves only through knowing
others. The genetic order, so far as there is an order, is, they
say, not first the ego and then the alter, but first the mother
and companions and then through them the self. Or, to put
this point of view in a less questionable form, it is only
through the reaction of one on the other that the two are
differentiated. Be this as it may, it is only through the action
of environment on the organism, and the reaction of the organ-
ism on the environment in behaviour, that experience becomes
polarized into subject and object. Let it be clearly understood
that for the animal, in all probability, subject and object are
not clearly distinguished, and set over against each other in the
antithesis of thought. Only late in mental development are
the self and the world distinguished in subtle analysis as
different aspects of the common experience in which both
have their inseparable being. Animals, and perhaps the
majority of mankind, never trouble themselves about object
and subject as clearly realized products of conception and
reflective thought. For these concepts are exceedingly subtle.
And here, too, the external aspect of experience has the pre-
cedence, so far as there is precedence. A healthy lad from
the moment he gets up in the morning till the moment he
goes to bed, lives chiefly in the objective aspect of experience,
an aspect which is in us chiefly associated with the products in
consciousness of the leading senses of sight and hearing.
But the subjective aspect creeps in when he is hurt, when he
is hungry, when he is fatigued. He does not argue about the
matter, or formulate it in definite terms. He just dimly

246 THE FEELINGS AND EMOTIONS

feels that the interest has somehow shifted. Still more dimly
does the animal feel that, apart from external interests which
prompt nine-tenths of its behaviour, chiefly through the senses
of sight, hearing, and smell, there are also matters in which
the interest has somehow shifted to his own body. For the
germ of self is essentially an embodied self. And perhaps
the emotions, which ring through the system for some time
after the external cause has been removed, serve in some degree
to aid in this dimly felt shifting of interest.

Whatever may be the exact psychological nature of the
emotions—and there has been much discussion of the question
—it may be regarded as certain that they introduce into the
conscious situation elements which contribute not a little to
the energy of behaviour. They are important conditions to
vigorous and sustained conation. And so closely interwoven
are these elements with the whole situation in its impulsive
aspect, that their disentanglement, in psychological analysis,
isa matter of extreme difficulty. I have elsewhere * devoted
some space to the consideration of the matter. I there follow
Professor William James in regarding organic effects, other
than motor sensations, as specially characteristic of the emotions
in their primary genesis. The cold sweat, the dry mouth, the
catch of the breath, the grip of the heart, the abdominal
sinking, the blood-tingle or blood-staguation—these and their
like, in varied modes and degrees, characterize the emotions
of fear, dread, anger, and so forth, when they rise to any pitch
of intensity, and contribute largely to their sharpness and
piquancy. These organic effects may be regarded as part of
the private and individual business of the body; but in ex-
perience they closely coalesce with the motor effects through
which the animal has to deal in practical behaviour with
that which evokes the emotion.

On this view these organic states which contribute cha-
racteristic elements in the emotional consciousness are due to
afferent, data from the vascular system and visceral organs,
just as motor consciousness is due to afferent data from the

* “ Habit and Instinct,” ch. ix., p. 186.

IMPULSE, INTEREST, AND EMOTION 247

parts concerned in overt behaviour. But, associated with
emotional states, there are also certain motor reactions, which
we speak of as their “‘ expression ”—so carefully discussed and
elucidated by Darwin,—and these unquestionably contribute
data to consciousness which coalesce with those afforded by
the visceral and vascular elements. The whole is commonly
suffused with feeling-tone, and the object which excites the
emotion is a centre of pleasurable or painful interest. Repre-
sentative elements, as experience develops, crowd into the
conscious situation and render it more complex. And in
addition to all this, there is, apart from the motor expression,
the strenuous behaviour of flight or attack, or other mode of
vigorous procedure which we commonly speak of as the
outcome of the emotional state. The conscious situation, in
the case of an enraged or scared animal actually behaving as
such, is thus exceedingly complex. And it should be under-
stood that in urging the importance of vascular and visceral
elements, this complexity is nowise denied. What is suggested
is that these elements are essential, and that they serve to
characterize the distinctively emotional factor in the situation,
that in any case they heighten the conative tendency.

Sufficient has now been said to indicate—but scarcely more
than indicate—the importance of feeling-tone, interest, and
emotion in determining the nature, character, and effective
energy of the conscious situations which arise in the course
of animal behaviour. They largely influence, and in part
direct, the course of the conative tendency. But they also
occur as its sequel. In animal,as in human life, the successful
attainment of the end towards which conation sets is highly
pleasurable. The equilibrium that is reached after instability,
though it marks the close of present endeavour, leaves after-
effects in consciousness in a sense of satisfaction which enters
re-presentatively into later situations and helps to further
more strenuous endeavour.

248 THE FEELINGS AND EMOTIONS

IJ.—Puay

“There are two quite different popular ideas of play,” says
Professor Groos, in his admirable work on “The Play of
Animals.” “The first is that the animal (or man) begins to
play when he feels particularly cheerful, healthy, and strong ;
the second that the play of young animals serves to fit them
for the tasks of later life.” The former view, in which the
latter may be included incidentally as a result, is closely asso-
ciated with the names of Schiller,* who suggested it, and of
Mr. Herbert Spencer,t who developed it. Mr. Wallaschek t
expresses the conception briefly and clearly when he says,
“It is the surplus vigour in more highly developed organisms,
exceeding what is required for immediate needs, in which play
of all kinds takes its rise, manifesting itself by way of imita-
tion or repetition of all those efforts and exertions which are
essential to the maintenance of life.” That surplus vigour is
often a condition favouring the manifestation of play is prob-
able enough, and seems to be supported by observation and
experience ; but that it is likewise a condition favouring the
chase, combat, mating, and much of the serious business of
animal life seems equally unquestionable. Success in all these
matters is largely determined by overflowing energy. In play,
however, this surplus vigour finds vent when there is no
serious occasion for its exercise. But, as Professor Groos says,§
“while simple overflow of energy explains quite well that the
individual who finds himself in a condition of overflowing
enerey is ready to do something, it does not explain how it
happens that all the individuals of a species manifest exactly
the specific kind of play expression which prevails with their
own species, but differs from every other.” And if to this it
be replied, that the specific kind is determined by repetition or

* “Netters on the Adsthetic Education of Mankind,” xxvii.
+ “Principles of Psychology,” § 533.

t “On the Origin of Music,” pp. 231, 232.

§ “The Play of Animals,” Eng. trans., p. 12.

PLAY 249

imitation of what we have called the serious business of animal
life, Professor Groos’s rejoinder is,* that “the conception of
imitation here set forth—namely, as the repetition of serious
activities to which the individual has himself become accus-
tomed—cannot be applied directly to the primary phenomena
of play—that is, to its first elementary manifestations” prior
to any experience of these serious activities. The repetition
(with a difference !) is in such cases not the re-enactment of
what has been previously performed in full earnest by the
individual, but rather the reappearance in the young of ances-
tral modes of procedure—in other words, its specific character
is such because it is a piece of instinctive behaviour or arises
from instinctive proclivities. And this is the central point of
the interpretation elaborated with great skill and candour by
Professor Groos. Play is instinctive ; and its biological value
lies in the training it affords for the subsequent earnest of
life.

Before leaving the surplus energy theory of play one more
point made by Professor Groos may be mentioned. He con-
tends that, though superabundant energy is a favouring con-
dition of animal play (as it is, indeed, of all animal behaviour),
still it is not a necessary condition. Animals often play when
they are tired out. ‘“ Notice a kitten when a piece of paper
blows past. Will not any observer confirm the statement that,
just as an old cat must be tired to death or else already filled
to satiety if it does not try to seize a mouse running near it, so
will the kitten, too, spring after the moving object, even if it
has been exercising for hours and its superfluous energies are
entirely disposed of ? Or observe the play of young dogs. when
two of them have raced about the garden until they are forced
to stop from sheer fatigue, and they lie on the ground panting,
with tongues hanging out. Now one of them gets up, glances
at his companion, and the irresistible power of his innate
longing for the fray seizes him again. He approaches the
other, sniffs lazily about him, and, though he is evidently only
half inclined to obey the powerful impulse, attempts to seize

* Op. cit., p. 7.

250 THE FEELINGS AND EMOTIONS

his leg. The one provoked yawns, and in a slow, tired kind of
way puts himself on the defensive ; but gradually instinct con-
quers fatigue on him too, and in a few minutes both are tear-
ing madly about in furious rivalry until want of breath puts an
end to the game. And so it goes on with endless repetition,
until we get the impression that the dog waits only long
enough to collect the needed strength, not till superfluous
energy urges him to activity.” *

Coming now to Professor Groos’s interpretation of play, we
find in it, perhaps for the first time in the literature of the
subject, adequate stress laid on its biological value. ‘ The
play of young animals,” he says,f ‘‘ has its origin in the fact
that certain very important instincts appear at a time when
the animal does not seriously need them. . . . Its utility con-
sists in the practice and exercise it affords for some of the
more important duties of life, inasmuch as selection [in the
higher animals] tends to weaken the blind force of instinct,
and aids more and more the development of independent intel-
ligence as a substitute for it. At the moment when intelli-
gence is sufficiently evolved to be more useful in the struggle
for existence than the most perfect instinct, then will selection
favour those individuals in whom the instincts in question
appear earlier and in less elaborated forms—in forms that are
merely for practice and exercise,—that is to say, it will favour
those animals which play. ... Animals cannot be said to
play because they are young and frolicsome, but rather they
have a period of youth in order to play; for only by so doing
can they supplement the insufficient hereditary endowment with
individual experience, in view of the coming tasks of life.”

Some stress is here laid on the fact that important instincts
appear at a time when the animal does not seriously need them.
It seems to imply the doctrine of what biologists term “ accele-
ration ’—which means the development of an organ or mode
of behaviour at an earlier period in the descendants than that
at which it appeared in the ancestors. Thus the adult fighting
or hunting instinct of past generations appears in the young

* Op. cit., p. 19. t Op. cit., pp. 75, 76.

PLAY 251

to-day as a fighting-play or a hunting-play. It is open to
question, however, whether either the instinctive behaviour or
the conscious situation in the one case and the other is so
nearly identical that the playful fight or hunt can fairly
be called the same instinctive procedure as the serious
combat or chase. We may hold, with Professor Groos, that
the one is an invaluable preparation for the other without
identifying them as the same behaviour under different con-
ditions. Indeed the conditions are so different that the identi-
fication seems strained. The question may be left open,
however, without impairing the value of Professor Groos’s
suggestion. And we may divide the preparatory behaviour in
what is commonly called play under two heads: first, general
preparation for varied modes of serious effort in after-life ; and
secondly, special preparation for particular forms of this after-
effort. Under the first will fall what Professor Groos terms
experimentation and movement play, including what Dr. Stout,
who fully realizes its importance, calls “manipulation ” ; *
under the second, such forms as hunting-play and fighting-
play.

Nothing is more characteristic of the young of intelligent
animals than the variety and persistency of their behaviour,
their sensitiveness to stimuli of many different kinds, their
restlessness of swiftly changing attention and response, with
occasional pauses of continued effort in some special direction.
Constantly on the alert, they exhibit in all its shifting phases
behaviour which we interpret as indicating curiosity, inquisitive-
ness, love of mischief, destructiveness, and so forth. The facts
are so familiar to every observer of young animals that it is
unnecessary to give any detailed illustration. Watch a kitten
in this stage of its development and carefully note its behaviour
during half an hour; the variety of effort, the réles played
by trial, failure, and success, the gain of skill and control over
behaviour, will at once be evident. Or devote an equal space
of time to observing young jays, magpies, or jackdaws. Every
projecting piece of wire or bit of wood in their cage is pulled

* “ Manual of Psychology,” p. 327.

252 THE FEELINGS AND EMOTIONS

at this way and that way, from above, from below, from the
side. Now one, then another, loose object is picked up and
dropped, turned over, carried about, pulled at, hammered at,
stuffed into this corner and into that, and experimented with
in all possible ways. Then the wise bird goes to sleep, and
wakes up again only to resume with new zest its persistent
and varied efforts, by which it becomes acquainted with all
the details of its environment. Watch young birds on the
wing gaining their mastery of the air in flight, young seals
tumbling in the water, young foals scampering and kicking
up their heels in the meadows. A little observation, as occasion
serves, a little attention to the progress towards an adequate
experience of the meaning of things, towards more com-
plete control, and increased nicety of behaviour, whether in
reference to their surroundings, or in powers of finished
locomotion, will serve to bring home what Professor Groos
includes under experimentation and movement plays. He
regards it all as play, since it seems to have no serious end,
and is just a preparation for the sterner realities of adult life.
And for human beings, whose work is so largely enforced,
the freedom and evident joy of it all suggests the play
which has acquired for us the meaning of relaxation from
irksome effort, and glad abandonment to less constrained
modes of behaviour. But in young animals such play is, after
all, the serious business of their time of life. Its import
for their future welfare can scarcely be overestimated.

And its import is in large degree psychological. If we
watch a young puppy or kitten learning gradually to deal
effectively with some difficulty in its extending environment,
we see that it puts forth its efforts at first in a somewhat
random and indefinite fashion. It is one of those animals
in which intelligence has been evolved to supersede and
become the more plastic substitute for instinct. The random
and indefinite movements, are in detail reflex responses to
stimuli. But whereas, in a piece of highly elaborated instinc-
tive behaviour, such reflexes are grouped into a whole which
is co-ordinated through inherited nervous mechanism ; in the

PLAY 253

case of the acts of the puppy or kitten they have to be further
co-ordinated, or more elaborately grouped, through experience.
To act in one way some of the reflexes have to be checked as
redundant and not to the point: to act in another way other
reflexes have to be similarly checked; and in a third way,
yet others. But in all three some of the reflexes are utilized
to different ends. Many conscious situations contain common
elements; and this tends to give unity to the developing
experience. But they contain also elements and groupings
which afford that diversity without which conscious behaviour
could not be accommodated to them. So that we have here
the conditions under which what is technically termed “the
concomitant differentiation and integration of experience”
can proceed.

And if we speak of the instinct of experimentation we
must remember that what we are dealing with is rather an
innate tendency or instinctive propensity than a definite and
relatively clean-cut piece of instinctive behaviour. It com-
prises a great number of inherited reflex acts, and may perhaps
be fairly called instinctive in detail. But experimentation
must be regarded rather as the proximate end of a conative
tendency, or group of conative tendencies, whose ultimate
biological end is success in dealing with the environment in
the sterner struggle for existence during adult life. The
tendency is inherited, and therefore falls under the head of
instinctive propensity. But “experimentation” is a group-
term under which we comprise the general drift of varied
modes of behaviour, founded indeed on a congenital basis,
but receiving its stamp and character from what is acquired
in the course of the experience it provides. It is essentially
a process whereby the conscious situations acquire what
Dr. Stout terms meaning; and is specially interesting as
affording an example of the way in which intelligence moulds
and refashions a number of disconnected reflex responses. And
if, following Professor Groos, we call it play, it is a little
difficult to see how it can be brought in line with his statement *

* Op. cit., pref., p. XX.

254 THE FEELINGS AND EMOTIONS

that “the play of youth depends on the fact that certain
instincts, especially useful in preserving the species, appear
before the animal seriously needs them.” Does experi-
mentation occur before it is needed in the economy of animal
behaviour? And might we not with equal truth say that the
play of youth depends on the fact that certain acquired habits,
especially useful in preserving the species, are gained before
the animal seriously needs them ?

Passing now to those forms of play which afford more
special preparation for particular forms of after-effort, under
which fall such types as hunting-play and fighting-play, we
‘may refer the reader to the copious examples so carefully
collected by Professor Groos. The way in which a kitten pats
a cork or a ball, making it roll and then pouncing upon it, is
a characteristic example of animal play. Valuable as a pre-
paration for dealing successfully with a mouse when occasion
shall arise, this is a specialized form of experimentation ; and
it is more obviously in line with the hunting-behaviour of
later life than is general experimentation with any particular
modes of future behaviour. Still it is essentially experimenta-
tion, with the instinctive propensity setting in more definite
channels. Its value lies in the acquisition of skill under
circumstances easier than those presented in the serious chase.
So, too, in the case of the playful tussles of puppies or in that
of the kitten, which not only shows playful fight to its
brothers and sisters, but also to its mother, who responds by
holding down the struggling and scratching little creature.
Unquestionably, there is an instinctive propensity ; much of
the detail, and some of the grouping, exhibit inherited reflexes
due to special modes of stimulation. No doubt many of these
responses occur in a similar but more emphatic way in a serious
fight, and yet we may hesitate before committing ourselves to
the theory of acceleration. It is at least equally probable
that play as preparatory behaviour differs in biological detail
(as it almost certainly does in emotional attributes) from the
earnest of after-life, and that it was evolved directly as a
preparation, as a means of experimentation through which

PLAY 255

certain essential modes of skill were acquired,—those animals
in which the preparatory play propensity was not inherited in
due force and requisite amount being worsted in the combats
of later life, and eliminated in the struggle for existence.
For, in the preparatory tussles and squabbles and playful fights
of young animals, experience is gained without serious risk to
life and limb.

The modifications of Professor Groos’s biological interpreta-
tion of play which we would suggest are so slight that we
may be said to accept it almost unreservedly. The play of
youth, we may urge, depends on instinctive propensities to
experimentation in varied ways, some of more general and
others of more special import ; and the value of such experi-
mentation lies in the fact that it is a means of acquiring,
under circumstances more easy and less dangerous than those
of sterner life, experience and skill for future use. In a word,
play depends on instinctive propensities of value in education.

Passing now to a brief consideration of the feelings and
emotions which we may suppose to accompany play, we may
place first those which characterize, from this point of view,
general experimentation. We have here rapidly varying
situations charged with conative impulse, the satisfaction of
which must bring pleasure—the occasional thwarting of which
is probably toned with the opposite—the latter serving, through
contrast, to enhance the satisfaction of ultimate success. Both
pleasure and its antithetical state of feeling are primarily
matters of the conscious situation as a whole, and even in
ourselves are difficult to distribute in analysis. But assuredly
no small share of the total product must be assigned to the
successful behaviour which consummates the conative tendency.
Indeed, it is the thwarting of free action which is the source
of much of the discomfort of the young. Unimpeded and
vigorous behaviour also brings with it secondary effects in organic
processes—fuller heart-beat, freer circulation, deeper respira-
tion, better digestion, firmer muscular tone, and so forth—
which have a marked effect on the conscious situation, and aid
in producing that emotional tone which cannot, perhaps, be

256 THE FEELINGS AND EMOTIONS

named in better terms than “good spirits” and the joy of
existence, so forcibly suggested during the free play of youth.
On the other hand, there is no more piteous sight than
that afforded by the young animal, “cabined, cribbed, con-
fined,” suffering from ennwi and depression—all its organic
processes sluggish and craving to be quickened into the
natural vigour of life, not creeping slowly through the veins,
but coursing at full flood.

In the psychological aspect of play Dr. Groos assigns
perhaps the first place to pleasure in the possession of power,
or, as Preyer phrases it, pleasure in being a cause. We must
be careful, however, lest in using such expressions we seem to
imply that animals—even quite young animals—are capable
of entertaining ideas which belong to a much later stage of
mental development. Speaking of “joy in ability or power,”
Professor Groos says,* “This feeling is first a conscious pre-
sentation to ourselves of our personality as it is emphasized
by play. . .. But it is more than this; it is also delight in
the control we have over our bodies and over external objects.
Experimentation in its simple as well as its more complicated
forms is, apart from its effect on physical development, educa-
tive in that it helps in the formation of causal associations.
... The young bear that plays in the water, the dog that
tears a paper into scraps, the ape that delights in producing
new and uncouth sounds, the sparrow that exercises its voice,
the parrot that smashes his feeding trough—all experience the
pleasure in energetic activity, which is, at the same time, joy
in being able to accomplish something.” But those who
agree with Dr. Stout, as E do without hesitation, in denying
personality (save in a very embryonic condition) and the con-
ception of causation to animals in the perceptual stage
of mental evolution, though they may find in Dr. Groos’s
contention a central core of truth, will be unable fully to
accept his manner of presenting it. ‘Any single train of
perceptual activity,” f says Dr. Stout, “has internal unity and

* Op. cit., p. 290.
t+ “Manual of Psychology,” p. 266.

PLAY 257

continuity. But where conscious life is mainly perceptual, the
several trains of activity are relatively isolated and discon-
nected with each other. They do not unite to form a
continuous system, such as is implied in the conception of a
person. We must deny personality to animals.” To this I
would merely add that, even where perceptual continuity in
animals reaches its maximum, it is not reflectively grasped as
a whole, and the ideal construction of the personal ego is not
conceived as antithetical to the impersonal world of objects.
With what Dr. Stout says about causality I am in complete
agreement. “We must notice,’* he urges, “the essential
difference which separates the merely perceptual category from
that of ideational and conceptual thought. The perceptual
category is always purely and immediately practical in its
operation. It is a constitutive form of thought only because
it is a constitutive form of action. The question ‘Why ?’ has
no existence for the merely perceptual consciousness. It does
not and cannot inquire how it is that a certain cause produces
a certain effect. It does not and cannot endeavour to explain,
to analyze conditions so as to present a caus? asa reason. It
does not compare different modes of procedure or different
groups of circumstances, so as to contradistinguish the precise
points in which they agree from those in which they disagree,
and in this way to explain why a certain result should follow
in one case and a different result in another case. Causality
in this sense can only exist for the ideational consciousness,
and the development of the ideational consciousness in this
direction is a development of conceptual thinking—of gene-
ralization.”

Wherein, then, lies the central core of truth in Professor
Groos’s contention? In the satisfaction that arises from the
success of any conative activity. We see that the animal
striving and doing falls within our conception of a cause, in
the scientific sense of the word,—a relatively constant and
continuous antecedent of diverse sequent effects. We infer
that pleasure accompanies the satisfaction of the multifarious

* Op. cit., p. 314.
8

258 THE FEELINGS AND EMOTIONS

conative impulses. The pleasure is the animal’s ; the conception
of causality and of self as a continuous person, still the same
amid diversity of conscious situations, is ours. If we bear this
in mind there can be no objection to our attributing to
animals joy in ability or power. It is the pleasure derived
from that successful conation whereby animals fall into the
category of causes within the scheme of our rational thought.

In fighting-play and hunting-play, too, there arise in
more specific forms the pleasures of successful conation with
the antithetical feelings accompanying thwarted conation.
And these are distinguished from earnest, partly because the
companion or the inanimate substitute for prey is the centre
of a different situation from that afforded by an enemy or the
natural object of the chase; partly by the absence of certain
insistent emotional states which characterize earnest and
the serious business of life. In fighting, this is anger. And
we often see the tendency of this to arise in the midst of
fighting-plays, and at once say that it becomes serious and
passes into fighting in earnest. Indeed, some tinge of earnest,
with its fuller emotional tone, forms part of the preparation
for future life, and so far falls within the definition Professor
Groos gives of play. From which we may see that play is not
easily marked off from other forms of conation.

Brief reference to the element of ‘ make-believe,” which
Dr. Groos assigns to the higher forms of play, may be reserved
for our fourth section; and some further discussion of its
psychological aspect to the concluding chapter.

III.—CovurtsHie

We have seen that Professor Groos regards play as the
practice and preparation for the serious business of animal
life. Founded on instinctive tendencies, it has its biological
value in the acquisition of practical acquaintance with the
environment, and of skill in dealing with it effectually. It is
an education in behaviour of the utmost service in view of the
struggle for existence, It is full of the pleasure derived from

COURTSHIP 259

the satisfaction of innate impulse, the success of conative
effort, and the diffused sense of well-being which accompanies
a life of action, free and unrestrained. This freedom and
gladness lead us to call it play ; but we must not draw the
inference that the playful animal knows that it is playing, or
forms any conception of the antithesis between work and play,
which is a product of late development.

In laying stress on the biological value of certain modes
of behaviour which we thus call play, a value which lies in the
practice and preparation they afford for life’s more earnest
work, Professor Groos deserves our hearty thanks. Nor need
our thanks be less hearty if we find that he has in some
degree been anticipated by Darwin; for he has elaborated
with systematic care what Darwin suggested incidentally.
“ Nothing is more common,” said Darwin,* “ than for animals
to take pleasure in practising whatever instinct they follow at
other times for some real good. How often do we see birds
which fly easily, gliding and sailing through the air, obviously
for pleasure! The cat plays with the captured mouse, and
the cormorant with the captured fish. The weaver bird, when
confined in a cage, amuses itself by neatly weaving blades of
grass between the wires of its cage. Birds which habitually
fight during the breeding season are generally ready to fight
at all times ; and the males of the capercailzie sometimes hold
their Balzen, or leks, at the usual place of assemblage during
the autumn. Hence it is not at all surprising that male birds
should continue singing for their own amusement after the
season of courtship is over.”

In the behaviour of courtship we have what is essentially
part of the serious business of animal life. And in including
it under the heading of ‘“ Love Plays,” Professor Groos may
seem to be forgetful of his own definition of play. He is,
however, too clear a thinker not to see, and too honest an
exponent not to say, that much of the emotional behaviour
commonly regarded as courtship falls outside his main thesis
“in being, strictly speaking, not mere practice preparatory to

* “ Descent of Man,” vol. ii., p. 60, 2nd edit. 1888.

260 THE FEELINGS AND EMOTIONS

the exercise of an instinct, but rather its actual working.” *
But behaviour of a somewhat similar kind is seen in young
animals before the time of mating has arrived, and is exempli-
fied both in young and adults under circumstances different
from those which distinguish what we may term the pairing
situation. This, at any rate, may be regarded as a form of
experimentation and practice in the arts of courtship. On
different grounds does Professor Groos attempt to justify the
inclusion of actual courtship under the head of play. For it
may also, he thinks, even at the time of ‘its serious exercise, be
to some extent artful, involving ‘“ make believe,” and there-
fore playful in a somewhat different and more subtle sense ;
but a brief reference to “make believe” we may reserve for
our next section.

There can be no question that special modes of behaviour
often characterize the pairing situation, and that these not
only exemplify an instinctive tendency, but from their con-
stancy and relative definiteness constitute types of instinctive
behaviour. They would form parts of any definition of a
species founded not on structure but on behaviour. And if
animals have feelings and emotions at all—if they are not
Cartesian automata, which merely seem to be guided in their
actions by consciousness—there can be but little question that
the behaviour which characterizes the sexual situation is un-
usually charged with feeling-tone, and accompanied by all
those adjuncts which distinguish an emotional state, broadly
considered. This matter is of no little importance in our
interpretation of the phenomena described as courtship. Do
the accompanying feeling-tone and the state of emotional
exaltation influence the behaviour, or would it run a similar
course in the absence of any such accompaniments? If, as we
can scarcely doubt, the consciousness attending the situation
does profoundly influence the behaviour, the further question
arises—Is this influence mainly the result of the presence and
behaviour of an individual of the oppositesex ? To this, again,
we must answer that, so far as we can learn by observation,

* “The Play of Animals,” Eng. trans., p. 229.

COURTSHIP 261

behaviour unquestionably is determined by such influence
in the serious business of courtship. And then the further
question arises—Is it a matter of indifference what the appear-
ance and behaviour of the individual of the opposite sex may
be? AreA.,B., C.,and the rest of the male alphabet, precisely
alike in stimulating in a similar manner, and to a similar
degree, the sexual impulse of the female? If we admit any
differential influence, and if this influence takes effect in the
sexual union to which courtship is preparatory, we so far admit
the efficacy of that which Darwin termed sexual selection.

Let us, however, before proceeding with general con-
siderations, present one or two examples of the facts which
observation has furnished with regard to specialized modes
of behaviour at the time of pairing. Speaking of American
night-hawks, Audubon says, “ Their manner of flying is a
good deal modified at the love season. The male employs
the most wonderful evolutions to give expression to his
feelings, conducting them with the greatest rapidity and
agility in sight of his chosen mate, or to put to rout a rival.
He often rises to a height of a hundred metres and more, and
his cries become louder and louder as he mounts; then he
plunges downward with a slanting direction, with wings half
open, and so rapidly: that it seems inevitable that he should be
dashed to pieces on the ground. But at the right moment,
sometimes when only a few inches from it, he spreads his wings
and tail, and, turning, soars upward once more.” * Mr. Strange,
quoted by Darwin,f says of the satin bower-bird, “at times the
male will chase the female all over the aviary, then go to the
bower, pick up a gay feather or a large leaf, utter a curious
kind of note, set all his feathers erect, run round the bower
and become so excited that his eyes appear ready to start from
his head ; he continues opening first one wing, and then the
other, uttering a low, whistling note, and, like the domestic
cock, seems to be picking up something from the ground, until
at last the female goes quietly towards him.”

* Quoted by Groos, op. cit., p. 259.
t “Descent of Man,” vol. ii., p. 77.

262 THE FEELINGS AND EMOTIONS

Darwin describes how, in the Argus pheasant,* ‘the im-
mensely developed secondary wing-feathers are confined to the
male, and each is ornamented with a row of from twenty to
twenty-three ocelli, each above an inch in diameter. These
beautiful ornaments are hidden until the male shows himself
before the female. He then erects his tail, and expands his
wing-feathers into a great, almost upright, circular fan or
shield, which is carried in front of the body. The ocelli are so
shaded that, as the Duke of Argyll remarks, they stand out
like balls lying loosely within sockets. But when I looked at
a specimen in the British Museum, which is mounted with the
wings expanded and trailing downward, I was,” adds Darwin,
“ oreatly disappointed, for the ocelli appeared flat or even con-
cave. Mr. Gould, however, soon made the case clear to me, for
he held the feathers erect, in the position in which they would
naturally be displayed, and now, from the light shining on
them from above, each ocellus at once resembled the ornament
called a ball and socket.” The primary wing-feathers are
scarcely, if at all, inferior in beauty to the secondaries, though
the markings are quite different, the chief ornament being a
space parallel to the dark-blue shaft, which in outline forms
a perfect second feather lying within the true feather. “Now
the secondary and primary wing-feathers are not at all dis-
played, and the ball and socket ornaments are not exhibited in
full perfection, until the male assumes the attitude of court-
ship.”

It is unnecessary to describe the song of birds which is
generally, but not always, at its best during the period of
pairing. Bechstein, who kept birds during his whole life, and
studied them with care, asserts that “the female canary always
chooses the best singer, and that in a state of nature the
female finch selects that male out of a hundred whose notes
please her most.”

Thus we are led back to sexual selection. If we are satisfied
that the males of certain species do as a matter of fact behave

* Darwin, op. cit., p. 99.
+ Quoted by Darwin, ‘ Descent of Man,” vol. ii., p. 58.

COURTSHIP 263

in specific and distinguishable ways at the breeding season, and
in presence of their would-be mates, the question is, What, if
any, is the biological value of such behaviour? What has
fostered and guided it in the course of its evolution? From
the case of the Argus pheasant, which is only a sample of the
large class of cases in which the male has special adornments,
we see that the behaviour has often direct relation to the
display of such plumage, or, in some apes, of coloured surfaces,
so that behaviour and ornamentation must be taken together.
The essence of Darwin’s contention is, that the adornments
and behaviour give rise to a situation through which the
female is stimulated or excited to accept the male; that the
male in which they are best developed vives rise to the most
effective situation, produces most excitement, and therefore
has the best chance of acceptance, being “ unconsciously pre-
ferred ;” * and that he thus begets offspring which inherit his
adornments and modes of behaviour, such inheritance being,
however, confined to the males. Thus sexual selection takes
effect through preferential mating, whereby certain hereditary
traits are transmitted and become racial characteristics. And
this is brought about through an appeal to consciousness, and
seems to involve choice—generally that of the female.

Now, as I have elsewhere urged,f the hypothesis of sexual
selection has often been placed in a false light by the introduc-
tion of the unnecessary supposition that the hen bird, for
example, must possess a standard or ideal of zesthetic value, and
that she selects that singer which comes nearest to her concep-
tion of what a songster should be. Darwin occasionally ex-
pressed himself unguardedly in the matter; he says, for
example, that the female appreciates the display of the male,
and places to her credit a taste for the beautiful.t But he also
distinctly states that “it is not probable that she consciously
deliberates ; she is most excited or attracted by the most
beautiful, or melodious, or gallant males.” § This is all that is
really necessary for the theory of sexual selection. The hen

* « Descent of Man,” vol. ii., p. 56. ¢ “Habit and Instinct,” p. 217.
t “Descent of Man,” vol. ii., p. 251. § Op. cit., p. 187.

264 THE FEELINGS AND EMOTIONS

accepts that mate which by his song or otherwise excites in
sufficient degree the pairing impulse ; if others fail to excite
this impulse, they are not accepted. Even Mr. A. R. Wallace,
who rejects the theory save in a very subordinate form, says i
that “it may be admitted, as highly probable, that the female
is pleased or excited by the display,” and speaks + of a possible
choice of “the most vigorous, defiant, and mettlesome male,”
giving, moreover,{ several telling examples of preferential
mating. Stripped of all its unnecessary wsthetic surplusage,
at any rate so far as this implies an esthetic ideal, or esthetic
motive, the hypothesis of sexual selection suggests that the
accepted mate is the one which adequately evokes the pairing
impulse.

Here Dr. Groos makes an interesting and important con-
tribution to the subject. He lays stress on the coyness and
reluctance of the female, and illustrates it by examples derived
from observation. “Thus the female cuckoo answers the call
of her mate with an alluring laugh that excites him to the
utmost, but it is long before she gives herself up to him.
A mad chase through tree-tops ensues, during which she con-
stantly incites him with that mocking call, till the poor fellow
is fairly driven crazy. The female kingfisher often torments
her devoted lover for half a day, coming and calling him, and
then taking to flight. But she never lets him out of her sight
the while, looking back as she flies and measuring her speed,
and wheeling back when he suddenly gives up the pursuit.
The bower-bird leads her mate a chase up and down their
skilfully built pleasure-house, and many other birds behave in
a similar way. The male must exercise all his arts before her
reluctance is overcome. She leads him on from limb to limb,
from tree to tree, until it seems that the tantalizing change
from allurement to resistance must include an element of
mischievous playfulness.” §

Professor Groos regards the instinctive coyness of the
female as the most efficient means of preventing the too early

* “Darwinism,” p. 285. t Op. cit. p. 293.
t Op. cit., p. 172. § Op. cit., pp. 285, 286.

COURTSHIP 265

and too frequent yielding to sexual impulse.* He thinks it
probable that “in order to preserve the species the discharge of
the sexual function must be rendered difficult, since the impulse
to it isso powerful that, without some such arrest, it might
easily become prejudicial to that end. This same strength of
impulse is,” he adds,f “itself necessary to the preservation of
the species ; but, on the other hand, dams must be opposed to
the impetuous stream, lest the impulse expend itself before it is
made effectual, or the mothers of the race be robbed of their
strength, to the detriment of their offspring.” It has its
origin in the general fact that, before any important motor
discharge takes place, there is apt to be a preparatory and
gradually increasing excitement. But this is specially empha-
sized in association with the sexual impulse. As Professor
Zeigler wrote, in a private communication to Dr. Groos,t
“ Among all animals a highly excited condition of the nervous
system is necessary for the act of pairing, and consequently we
find an exciting playful prelude very generally indulged in.”

Courtship may thus be regarded from the physiological
point of view as a means of producing the requisite amount of
pairing-hunger ; of stimulating the whole system and facilitating
general and special vascular changes ; of creating that state of
profound and explosive instability which has for its psycho-
logical concomitant or antecedent an imperious and irresistible
craving. This not only overcomes the coyness of the female,
but generates and strengthens the ardour of the male—a point
on which, perhaps, Professor Groos does not lay sufficient
stress. For the process is reciprocal ; and though the male
leads in the ardour of courtship, yet this ardour constantly
grows till at last it overcomes the barriers of reluctance.
Courtship is thus the strong and steady bending of the bow
that the arrow may find its mark in a biological end of the
utmost importance in the survival of a healthy and vigorous
race.

The coyness and reluctance of the female afford the

* Op. cit. p. 283. t+ Op. cit., p. 248.
t Op. cit., p. 242.

266 THE FEELINGS AND EMOTIONS

conditions under which the bow is bent to the full. But they
also afford the conditions of the apparent act of choice. This
takes the place, on the perceptual plane of mental development,
of that deliberation which precedes the higher act of choice on
the ideational plane. For psychology, as well as for biology,
then, Dr. Groos’s suggestion is a welcome and helpful one.
Both upholders of sexual selection and critics of that hypo-
thesis, have been apt to regard the choice of a mate in animals
from too anthropomorphic a point of view—to look upon it
as the outcome of rational deliberation, of weighing in the
esthetic balance the relative attractiveness of this suitor and
of that, and of reaching a definite conclusion that the one is to
be accepted because he behaves or is adorned in such and such
a way, while the other is to be rejected because he falls below
all reasonable standards of requirement. The choice exercised -
by the female, if so we term it, is far simpler and more naive.
Indeed, Professor Groos goes so far as to say that on his
hypothesis the element of choice is altogether abolished. “ It
is the instinctive coyness of the female,” he says,* “that
necessitates all the arts of courtship, and the probability is that
seldom or never does the female exert any choice. She is not
awarder of the prize, but rather a hunted creature. So, just
as the beast of prey has special instincts for finding his prey,
the ardent male must have special instincts for subduing
female reluctance. According to this theory, there is choice
only in the sense that the hare finally succumbs to the best
hound, which is as much as to say that the phenomena of
courtship are referred at once to natural selection.” He
reaches this conclusion, however, by gradual stages. He first
urges t that “it would be absurd to affirm that all bird songs
originate in a conscious esthetic and critical act of judgment
on the part of the female. A conscious choice either of the
most beautiful or the loudest singer is certainly not the rule,
and probably never occurs at all. But,” he adds, “is it not
still a choice, though unconscious, when the female turns to
the singer whose voice, whether from strength or modulation,
* Op. cit., pref., p. xxii. t Op. ctt., p. 240.

COURTSHIP 267

proves most attractive? Even if the song is primarily a
means of recognition or an invitation from the male, still the
psychological effect must be that the female follows the songster
that excites her most, and so exerts a kind of unconscious
selection.”

The phrase “ unconscious choice” is, however, somewhat
unsatisfactory, especially when we remember that it is used to
indicate the result of a direct appeal to the conscious situation.
If, however, we say that it is perceptual choice arising from
impulse as distinguished from ideational choice due to motive
and volition, we see that the distinction is in line with that
which we have drawn again and again, and which Dr, Stout so
well emphasizes in his “‘ Manual of Psychology.” But we have
also drawn the distinction between instinctive behaviour prior
to individual experience, and intelligent behaviour the result of
such experience. Under which of these classes does the be-
haviour of the female during courtship fall? Professor Groos,
in the further development of his hypothesis, seems to place it
in the instinctive category. ‘Instead of a conscious or un-
conscious choice, of which we know nothing certain,” he says,*
“we have the need of overcoming instinctive coyness in the
female, a fact familiar enough, but hitherto not sufficiently
accounted for. Then the question no longer is, which among
many males will be chosen by the female, but which one has
the qualities that can overcome the reluctance of the female
whom he woes. Sexual selection would then become a special
case of natural selection.”

I ain unable to follow Professor Groos in this view, which
I find it rather difficult to reconcile with such statements
as that already quoted, in which he says that the female’s
“tantalizing change from allurement to resistance seems to
include an element of a mischievous playfulness.” It is more
probable that instinctive coyness and reluctance afford the
conditions under which experience of the pleasures of courtship
may be gained. It is said that a flirt, when taken to task for
her conduct at ball and picnic, justified it by asking demurely

* Op. cit., p. 244.

268 THE FEELINGS AND EMOTIONS

how else she was to gain that wide experience of men which
was absolutely necessary to guide her in exercising a wise and
becoming choice. Let us hope that when the fateful time
arrived she acted with due deliberation. Now the coquetry
of birds affords the opportunity of gaining just such experience
in the light of which a perceptual choice may be made.

Let us remember that courtship is, as Darwin said, “a
prolonged affair,” and that coyness is a means to its prolonga-
tion. And let us remember that in simple cases, as also in
more complex matters, the intelligent exercise of choice
depends upon what Dr. Stout terms the acquisition of meaning.
“The chicken does not, at first,” he says,* “ distinguish
between what is edible and what is not. This it has to learn
by experience. It will at the outset peck at and seize all worms
and caterpillars indiscriminately. There is a particular cater-
pillar called the cinnabar caterpillar. When this is first
presented to the chicken it is pecked at and seized, like other
similar objects. But as soon as it is fairly seized it is dropped
in disgust. When next the chicken sees the caterpillar, it
looks at it suspiciously, and .refrains from pecking. Now,
what has happened in this case? The sight of the cinnabar
caterpillar re-excites the total disposition left behind by the
previous experience of pecking at it, seizing it, and ejecting
it in disgust. Thus the effect of these experiences [what I
have termed the conscious situation] is revived. The sight
of the cinnabar caterpillar has acquired a meaning.” Take
now the case of a coy hen bird, to whom several males pay
court. The sight of this one, behaving after his kind, excites
in small degree the sexual impulse and emotions. Her heart
beats but little the faster for all his antics, her respiratory
rhythin is scarcely affected, her feathers, like her feelings,
remain comparatively unrufled. He has acquired meaning
from the reaction to his presence ; it is not, however, a very
attractive meaning. But that other, perhaps from mere
persistency, perhaps because he is more “ vigorous, defiant, and
mettlesome” (she, at any rate, certainly knows not why), deeply

* “Manual of Psychology,” p. 85.

COURTSHIP 269

stirs her organic being, sets her all aglow, and breaks down the
barriers of her coyness. And this he does because he is the
centre of a conscious situation which has acquired, through
her experience of his presence, a meaning and an interest that
are at last irresistibly attractive. It is a choice from impulse,
not the result of deliberation; but it is a choice which is
determined by the emotional meaning of the conscious situation.
And it is the reiterated revival of the associated emotional
elements which generates an impulse sufficiently strong to
overcome her instinctive coyness and reluctance.

And this coyness is the natural correlative of the ardour
of the male, an ardour increased by his courtship antics. If
the female yielded readily and at once, the behaviour of
courtship would never have been evolved. Superabundant
vigour in the male is, no doubt, a favourable condition of
courtship, as it is of play ; but neither is it a sine gad non, nor
in any case does it, or can it, afford any guidance of behaviour
into just those specific channels in which we find it setting
during the breeding season. If sexual selection be not a vera
causa of the specific direction, we have at present no other
hypothesis which in any degree fits the facts. And to the
criticism of those who, like Mr. W. H. Hudson, urge that
dance and song, and aérial evolutions in birds, are seen at
times when the immediate business of courtship forms no part
of the situation, Professor Groos’s theory of play affords a
sufficient answer. If courtship, whose biological end is of
such supreme importance, forms a central feature in the serious
business of animal life; and if play is the preparation and
practice for behaviour of biological importance; we should
expect to find manifestations (with an emotional difference,
and no doubt many differences in detail) of all those actions,
the due performance of which, in the supreme hour of court-
ship, will alone enable the adequately prepared and well-
practised male to overcome the reluctance of the female, and
beget offspring to transmit his instinctive and emotional
tendencies.

4

270 THE FEELINGS AND EMOTIONS

IV.—AnimaL “ ASstHetics ” anp “ ETHICS”

In this section we shall consider some types of behaviour
which suggest situations that contain the germs of esthetics
and ethics, with a view to determining, so far as possible, the
principles on which they should be interpreted. This is a
peculiarly difficult subject; for we are endeavouring to get
behind the behaviour, and to infer the mental conditions which
accompany it, and through which it assumes its distinctive
character. The difficulty is twofold: first, because, as Dr.
Stout puts it,* “human language is especially constructed to
describe the mental states of haman beings, and this means
that it is especially constructed so as to mislead us when we
attempt to describe the workings of minds that differ in any
great degree from the human ;” and secondly, because, to
quote the same careful thinker,t “the besetting snare of the
psychologist is the tendency to assume that an act or attitude
which in himself would be the natural manifestation of a
certain mental process must, therefore, have the same meaning
in the case of another. The fallacy lies in taking this or that
isolated action apart from the totality of conditions under
which it appears. It is particularly seductive when the animal
mind is the subject of inquiry.”

We must, therefore, base our method of procedure on some
definite principle. The canon of interpretation which I have
elsewhere suggested { is, that we should not interpret animal
behaviour as the outcome of higher mental processes, if it can
be fairly explained as due to the operation of those which stand
lower in the psychological scale of development. To this it
may be added—lest the range of the principle be misunderstood
—that the canon by no means excludes the interpretation of a
particular act as the outcome of the higher mental processes,
if we already have independent evidence of their occurrence in

* “Manual of Psychology,’ p. 23.
t Op. cit., p. 22.
} “Introduction to Comparative Psychology,” p. 53.

ANIMAL “AESTHETICS” AND “ETHICS” 271

the agent. Now, the conclusion to which we are led by direct
experiment and a critical study of the actions of animals whose
life-history is known to us is, that most of their behaviour
—perhaps all—is due to what Dr. Stout terms the perceptual,
as opposed to the ideational, exercise of cognition. Their
behaviour can be explained without having recourse to the
hypothesis that they reflect, and attain to ideal schemes as the
result of abstraction and generalization consciously directed to
this end. Rather than repeat what I have already said, I will
quote Dr. Stout’s summary of the position to which he, too, has
been led. ‘The vast interval,’ he says,* “which separates
human achievements, so far as they depend on human intelli-
gence, from animal achievements, so far as they depend on
animal intelligence, is connected with the distinction between
perceptual and ideational process. Animal activities are either
purely perceptual, or, in so far as they involve ideas, these ideas
serve only to prompt and guide an action in its actual execu-
tion. On the other hand, man constructs ‘in his head,’ by
means of trains of ideas, schemes of action before he begins to
carry them out. He is thus capable of overcoming difficulties
in advance. He can cross a bridge before he comes to it.”

It has already been stated that in the intelligent behaviour
of animals under man’s teaching he is the rational agent, they
his willing slaves. This may be here again illustrated to
enforce the distinction drawn by Dr. Stout in the above
passage. Those who have seen a shepherd’s dog working
sheep on a moorland fell, and have taken the trouble to
ascertain how the results he sees have been attained, will
appreciate, on the one hand, how well the dog knows and
responds to the signals of his master, and, on the other hand,
how completely all initiation is in the master’s mind, not that
of the keenly intelligent dog. Those who merely witness such
a performance without inquiry or investigation will probably
misunderstand the whole matter. In the north of England
competitions are not uncommon where, say, three sheep have
to be driven over a definite course, between certain posts and

* “Manual of Psychology,” p. 266.

272 THE FEELINGS AND EMOTIONS

round others, through narrow passages and into a fold—all
within a certain time limit. At such a competition success
depends on two things: first, the training of the dog to re-
spond at once to some six or eight whistle-signals, often
accompanied by gestures and movements of a stick; and
secondly, the judgment of the shepherd. The signals, given
in different whistle-tones and inflections, have for the dog
meaning, such as drive straight on, from this side, from that,
stop, lie down, creep, and so forth. The dog’s whole business
is to obey these signals. And the instant response of a well-
trained dog is admirable. But in the whole proceeding he is
merely the executant of his master’s orders. He originates no
important step. And if you listen to the criticisms by other
shepherds during a competition you will find that they are
mainly passed on the judgment shown by the master, and
only in palpable failures in obedience on the behaviour of the
dog. The intelligent animal is what he is trained to be—one
whose natural powers are under the complete control of his
master with whom the whole plan of action lies.

Since, then, in the cognitional field we find no independent
evidence of the higher processes, we are bound, in accordance
with our canon, to interpret emotional situations on similar
principles, unless we find in them outstanding facts which can-
not be explained in this way.

In considering the pairing situation we urged that the
framing of an ideal of beauty to which a given suitor ap-
proaches, or from which he falls short, is unnecessary for the
interpretation of the facts. We should not in strictness, there-
fore, speak of ‘‘an appreciation of beauty ” or “a taste for the
beautiful” in birds, since such expressions almost inevitably
imply that these creatures have reached some conception of
beauty as distinguished from and contrasted with ugliness. At
the same time the hen certainly appears to enjoy the situation
of which the plumed cock, attitudinizing thus, forms the centre
of interest—through which he acquires meaning. Although,
therefore, there is probably no ideal or standard of beauty,
there are afforded the data in experience from which, were the

ANIMAL “ASTHETICS” AND “ETHICS” 273

bird capable of reflection, such an ideal might, in ideational
sublimation, be derived. Before comparison, abstraction, and
generalization can be applied, in the reflective laboratory of
thought, there must be suitable experiences to form the raw
material on which these rational processes can be exercised.
Long ere, in the course of mental evolution, the correlative
conceptions implied in the phrase “ beautiful or ugly” had
taken definite form, perceptual situations must have arisen,
where, by direct appeal to the senses, by the diffused effects of
stimulation and their accompanying feeling-tone, and by the
natural satisfaction of mere impulse, the foundations were laid
of that appreciation of the beautiful which forms the reflective
superstructure we build upon them. Indeed, the pleasure and
satisfaction attending particular situations, as they severally
arise, appear to contain the perceptual germs of what in later
development becomes aesthetic appreciation.

The bird which, having completed its nest, eyes it with
apparent satisfaction, may well have the germs of that which,
when rendered schematic in our thought, we call taste. Dr.
Gould, indeed, states that certain humming-birds decorate
' their nests “ with the utmost taste,” weaving into their struc-
ture beautiful pieces of lichen. And the gardener bower-bird
collects in front of its bower flowers and fruits of bright and
varied colours. What meaning these carry in the conscious
situation we do not know; we can only suppose that they
incidentally contribute to the heightening of the sexual impulse,
and have been evolved as a means of stimulation to the bio-
logical end towards which sexual selection is unconsciously
directed. For it is probable that all the situations with which
pleasure and satisfaction are in high degree associated are, in
primary origin, closely connected with behaviour directed,
through natural or sexual selection, to some definite biological
end, or, in brief, with behaviour of biological value. And it is,
perhaps, not improbable that the states of consciousness most
highly toned with strong emotion have their origin in those
situations which arise amid the pairing, parental, and com-
paniable relations of animal life.

T

274 THE FEELINGS AND EMOTIONS

We have already said that the companion, as the nucleus
of a situation, is a thing which reacts in altogether special ways,
so that it becomes differentiated from other things as some-
thing the meaning of which, and the interest in which, are sw
generis and unique intype. It becomes the centre of emotional
situations, which we ascribe to rivalry, emulation, jealousy,
and so forth. And we have also drawn attention to the view
that the genetic order, so far as there is an order, is not first
the ego and then the alter, but first the mother and com-
panions and then through them the self. We learn to know
ourselves only through knowing others. We must now ask
the question—a question which must be answered before we
can touch on the possible ethics of animals—how far, and in
what sense, the social animal regards others as of like nature
to itself, and capable also of like feelings and emotions.
Stated in this form we must, I think, answer the question in
the negative. The expression, “of like nature to itself,”
impligs that the self has already taken more or less definite
form, and that the animal infers that, since the alter behaves
and reacts in like manner to the ego, it also is an ego. This
is distinctly an act of reasoning. As Clifford phrased it, the
companion becomes an eject. We can never by direct ex-
perience become acquainted with the feelings of others, but
we can endow them ejectively with personality analogous to
our own.

But, though it is exceedingly doubtful whether any animal
can regard its companion as an “ eject,” may there not be a
perceptual anticipation of the ideational process that comes
with later-developed reflection? A decade ago I gave the
following answer to this question: “For myself, I cannot
doubt that animals project into each other the shadows of the
feelings of which they are themselves conscious.” * Professor
Mark Baldwin speaks of the stage at which this takes place,
as the “projective stage” of development. “Now, in the
fact,” he says,t “of herding, common life and arrangements

* “ Animal Life and Intelligence,” p. 340.
+ “Mutual Development in the Child and the Race,” p. 19.

ANIMAL “ASSTHETICS” AND “ETHICS” 275

for the protection of the herd, animal societies of various kinds,
animal division of labour, etc.,—whatever be the origin of it,—
we have what seems to be such an epoch in animal life. These
creatures show a real recognition of one individual by another,
and a real community of life and reaction, which is quite
different from the individualism of purely sensational and
unsocial consciousness. And yet it is just as different from
the reflective organization of human society, in which the
self-consciousness and personal volition of the individual play
the most important ré/e. I see no way of accounting for the
gregarious instinct anywhere, except on the assumption of such
a projective epoch of animal consciousness.”

Now, in endeavouring to realize how the situation feels to
an animal in this projective stage, the first difficulty we
encounter is that of divesting ourselves of those products of
reflection which characterize our own mental situation ; and
to avoid what Dr. Stout, in the passage above quoted,* terms
the psychologist’s besetting snare. The second difficulty is
to grasp that, in experience, subject and object are inseparable,
however clearly we may learn to perceive that they are dis-
tinguishable aspects of that experience. If the subject is
eventually regarded as that which experiences, and the object
as that which is experienced, it is surely obvious that each is
necessary to the other. But, before these different aspects are
clearly distinguished, there is, in the perceptual stage of
mental development, what we may term a distribution of the
items of experience among the centres of interest.

In illustration of the kind of distribution which we may
suppose to come naturally to an animal, in what Professor
Baldwin terms the projective epoch, let us take three animal
situations: first, a chick pecks at a soldier-beetle, and finds it
nauseous ; secondly, a hen-bird hears the joyous song of her
mate; thirdly, a puppy in play bites its companion, and
receives a painful nip in return. Each of these constitutes
an experience-situation ; assuming that the results of the expe-
rience are distributed, how may we suppose them to he allocated ?

* Supra, p. 270.

276 THE FEELINGS AND EMOTIONS

In the first case, the soldier-beetle is the centre of interest
in the situation. As the situation develops, the element of
nauseousness is introduced. As Dr. Stout puts it, this is what
gives the soldier-beetle meaning. Can it be doubted that,
if there be any distribution, the nauseousness, though it is
altogether what we have learnt to call a subjective affection,
attaches itself to the soldier-beetle? The plain man, un-
sophisticated by Berkeleyan discussion, says simply, in such
cases, “ The thing is nauseous.” And this probably indicates
the naive and primitive distribution. Turning now to our second
example, when the hen hears the courtship song the mate is the
centre of a situation suffused with pleasurable feeling. How
is the joyousness, again essentially subjective for our later
thought, distributed ? Surely, if at all, on the mate who forms
the centre of interest. This it is which gives him meaning.
The joy of the hearer is projected on to the singer. Not entirely,
perhaps ; the hen literally, on Professor James’s theory of the
emotions, feels her heart-beat quickened by his presence,
and the delightful ruffling of her feathers. But our aim is
not to deny that the germs of the subjective arise in the
midst of such situations, but to contend that some at least of
the joyous character of the situation attaches to the song
of the singer, that some of the feeling is projected, and that
this is what gives the mate meaning. In our third case, the
playful puppy bites his companion, and is sharply bitten in
return. Pain enters into the coalescent situation as a whole.
How is it distributed ? In the phraseology of association, the
nip he gives is closely linked with the pain he receives. By
coalescence the pain and the nip form parts of the developed
situation. But the companion is the centre of interest. And
part of the pain is probably projected on this centre. That
such projection actually occurs is rendered probable by such
cases as the following, which was told me some years ago. A
child, whose exact age I have forgotten if I then ascertained,
was pricked by a pin, and he said, ‘Pin ’urted; poor pin.”
It is, indeed, not unlikely that with animals the outward pro-
jection of feeling is widely distributed over inanimate, as well

ANIMAL “ ASSTHETICS” AND “ETHICS” 277

as animate, objects, and that its due restriction comes far later
in development, of which the so-called personification of life-
less things by savages may be a relic. In any case, the give-
and-take of play in young animals, and the after-earnest of
courtship and fighting, would seem to afford ample opportunity
for the external and internal distribution of feeling which
sows the seed in perceptual life of that which blossoms into
self and alter in the reflective life of ideational thought.
Although, therefore, an animal cannot conceive its com-
panion as another self of similar nature, and with like passions
to his own, yet a considerable share of the feeling-element of
the conscious situation is projected on to that companion as
the chief centre of interest. And if it be said that this is
his feeling and not his neighbour’s, the objection will be seen
to lose its force, so soon as it is realized that even man has
no experience of any feelings save his own. The only way we
can reach fellow-feeling is through sympathy ; and sympathy
has its roots in the projective process we have endeavoured to
describe. We endow our neighbours with natures as sensitive
to pain and pleasure as our own. This is a pre-requisite to
the social relationships termed ethical. But when we hear
people say, and find even Mr. Romanes putting on deliberate
record,* that “the feelings which prompt a cat to torture a
captured mouse can only be assigned to the category to which,
by common consent, they are ascribed—delight in torturing
for torture’s sake,” I venture to think that common consent,
if such it be, is wrong. As I said a dozen years ago,{ before
Professor Groos had so carefully elaborated his theory of play,
“the cat or kitten plays with the mouse not from innate cruelty,
but for the sake of getting some little practice in the most
important business of cat life. Only man, who has the
capacity for nobler things, can be cruel for cruelty’s sake ;”
and this is the direction in which Dr. Groos’s opinion { tends
to set. Mr. Romanes might have learnt a lesson in caution
* “ Animal Intelligence,” p. 413.

+ Atalanta, Jan., 1889. Reprinted in “ Animal Sketches,” p. 17.
+ “The Play of Animals,” p. 122.

278 THE FEELINGS AND EMOTIONS

from his sister, who at first attributed a sense of shame to the
capuchin she so carefully studied, but subsequently was led
to adopt a simpler interpretation. ‘He bit me in several
places to-day,” she says, in her admirable diary,* “but he
seemed ashamed of himself afterwards, hiding his face in his
arms, and sitting quiet for a time.” She adds, however, in a
footnote : “On subsequent observation, I find this quietness
was not duc to shame at having bit me; for whether he
succeeds in biting any person or not, he always sits quiet
and dull-looking after a fit of passion, being, I think,
fatigued.”

Shame is an ethical feeling. And as we have briefly dis-
cussed the germs of esthetics in animals, so we may now as
briefly consider the germs of ethics. In its developed form
ethics is one of the “ normative sciences” involving standards
of right and wrong. It is,as Professor Mackenzie says,t “the
science of the ideal in conduct.” It involves a standard of
“ought,” the product of reflection and generalization. Conduct
is compared with the ideal, and perceived to be either below,
up to, or perhaps beyond, the normal standard accepted by
civilized mankind. This involves a judgment; and so far
as conduct is shaped in accordance with the ideal we attribute
the guidance to ethical motives. Such ideals, such judgments,
and the control of conduct through the play of such motives,
are probably beyond the mental capacities of animals. They
belong to the ideational stage of mental development, when
the conative tendency becomes volitional; not to the per-
ceptual stage, when it is impulsive. They do not enter into
the conscious situation as it takes form in the animal mind.
Behaviour has not in them acquired ethical meaning, since
in developed ethics, as normative, such meaning always has
reference to the norm, or standard. A real sense of shame
implies that our acts have fallen below our ideal.

It may be said that we cannot prove that animals do not
frame such ideals. But, if we accept the canon of interpretation

* Appendix to “ Animal Intelligence,’’ p. 486.
tT “ Manual of Ethics,” p. 1.

ANIMAL “ ASSTHETICS” AND “ETHICS” 279

above laid down, what has to be proved is that they do
frame them. Is there any case among the hundreds that are
popularly adduced to show that dogs are ashamed of themselves,
that they possess a sense of justice, that they feel the prick of
conscience, that on the one hand they know when they have
done wrong, or on the other hand enjoy a sense of conscious
rectitude—is there any particular case so described in the
popular phraseology of anecdote, which could not be more
simply described ag the direct outcome of the coalescent
situation, without the introduction of any implied reference
to a standard of behaviour reached by reflective thought ?
The pug that has taken a nap on the drawing-room sofa, leaps
down and slinks off with a “guilty” look on his master’s
approach. One can surely picture the previous situations, and
be tolerably certain that they contained an element of reproof
or something more energetic. The poodle that has successfully
performed his tricks bounds to his mistress with an air of
duty well performed. Has he never been petted and patted
under such circumstance? Routine in many animals—so
often creatures of habit—begets a customary sequence, the
breach of which is at once felt. To this I ventured* to
ascribe the conduct of the turnspit dog reported by Arago.
He refused with bared teeth to enter out of his turn the drum
by which the spit was rotated. The companion dog was put
in for a few moments and then released ; whereupon the dog
which before had been so refractory seemed satisfied that his
turn for drudgery had come, and, entering the wheel of his
own accord, began turning the spit as usual. The bared teeth
may be here perhaps ascribed to an outraged sense of justice.
But is it not a more simple, and just as probable, supposition
that the behaviour was due to breach of customary routine.
A trainer with whom I had some conversation on this matter
pointed out a collie bitch, and said, “If I put her through her
tricks in the usual order she does them like an angel; but if
I try and make her alter the order she snaps and sulks like
the devil.”
* “ Animal Life and Intelligence,” p. 404.

280 THE FEELINGS AND EMOTIONS

I have elsewhere* expressed my opinion that, though
animals may behave in ways which may tend to mislead us,
they do not act with intent to deceive. A dog is described f
as “showing a deliberate design of deceiving” because he
hobbled about the room as if lame and suffering from pain
in his foot. But may not this be simply due to the fact that
chance experience had led to a situation through which a
hobbling gait had acquired the meaning of more petting
and attention than usual? To behave with deceit as a
deliberate motive implies the idea that the action will be
interpreted as having a significance different from that which
it really has. It is only possible on the ideational plane of
mental development. It implies, too, from the ethical stand-
point, a conscious departure from the standard of truth. The
black that is acted has conscious reference and relation to the
white that is not black. Few, however, will credit animals
with deceit of this fully conscious and deliberate kind. Like
the fibs of little children, the apparent deceit of animals is
probably merely behaviour which has been associated in
experience with pleasant results.

The case of shamming sickness, quoted from K. Russ,
is thus interpreted by Professor Groos.t And yet he adds,
‘““When we see deception used so effectively to serve practical
ends, examples of which are very common, it can hardly be
doubted that there is in all probability more consciousness of
shamming in play than we have any means of demonstrating.”
And elsewhere in the same work he observes,§ “ Many a
grown animal still takes pleasure in the mock combats that
he learned in youth. From a psychological point of view
this phenomenon is especially noteworthy, from the fact that
the adult animal, though already well acquainted with real
fighting, still knows how to keep within the bounds of play,
and must therefore be consciously playing a réle, making

* « Animal Life and Intelligence,” p. 400. “ Introduction to Com-
parative Psychology,” p. 369.

+ “ Animal Intelligence,” p. 444,

{ “The Play of Animals,” p. 299. § Op. cit., p. 145.

ANIMAL “ ASTHETICS” AND “ETHICS” 281

believe.” I fail, however, to see the justification for the
“therefore.” Surely the difference of behaviour in this
example, and in other such examples, is sufficiently explained
as the outcome of diverse situations, without having recourse
to anything so psychologically complex as the conscious self-
illusion of make-believe—interesting and important as this is
in the psychology of children. To suppose that a monkey
who nurses a bit of blanket has any ideas about its being a
make-believe baby is not to interpret the behaviour of animals
in accordance with the canon we have adopted for our
guidance.

To return to the “ethics” of animals. I have urged that
ethical ideas, properly so called, have no place in their psy-
chology. But just as the pleasure and satisfaction attending
particular situations, as they severally arise, appear to contain
the perceptual germs of what in later development becomes
esthetic appreciation; so, too, do they also contain the
perceptual germs of what becomes, through reflection in man,
ethical approbation. And the situations in which these ethical
germs must be sought are those which entail behaviour for
the good of the social community. Indeed, we may go so
far as to say that the perceptual foundations of ethics are laid
in the social instincts. The satisfaction or dissatisfaction
arising from the performance or non-performance of instinctive
behaviour, evolved for the biological end of the preservation
of the social community, is the perceptual embryo from which
conscience is developed. Professor Mackenzie has indicated
the ambiguities in the use of the term,“ conscience.” ‘ It is,”
he says,* “sometimes used to express the fundamental principles
on which the moral judgment rests ; at other times it expresses
the principles adopted by a particular individual; at other
times it means ‘a particular kind of pleasure or pain felt
in perceiving our own conformity or nonconformity to
principle. t+ This last seems to me,” adds Mr. Mackenzie,

* « Manual of Ethics,” pp. 285, 286.

+ Starcke, International Journal of Ethics, vol. ii., no. 3 (April, 1892),
p. 348,

282 THE FEELINGS AND EMOTIONS

“the most convenient acceptation of the term, except that
I should prefer to say simply that it is a feeling of pain
accompanying and resulting from our nonconformity to
principle.” According to this definition the existence of a
principle or ideal is presupposed ; and the fact that Professor
Mackenzie lays stress upon the pain of nonconformity, shows
that the ideal is a high one. In the case of the animal,
however, such an ideal of right conduct has probably not
taken form. But Mr. Mackenzie also speaks of the “ quasi-
conscience” begotten of custom. This comes nearer to the
feeling which animals may be supposed to have when their
behaviour does not accord with that which through instinct
or habit is the usage of the community. And if, as seems
to be shown by observation, animals sometimes punish the
breaches of such usage—when, for example, cats punish their
kittens for uncleanliness—the quasi-conscience will assume a
more developed form.

We may say, then, that the perceptual data are given in
animal experience from which, in ideational sublimation, ethical
ideals may be derived by a process of reflection and generaliza-
tion. As in the case of esthetics, so in that of ethics ; long ere,
in the course of mental evolution, the correlative conceptions
implied in the phrase “right or wrong” had taken definite
form, perceptual situations must have arisen in which
behaviour carried with it the feelings of satisfaction or the
reverse which laid the foundations of that approbation of the
right which forms the superstructure we build upon them by
the exercise of reflective thought.

V.—TueE Evo.ution or FEELING AND Emotion

“Whatever conditions,” says Dr. Stout,* “further and
favour conation in the attainment of its end, yield pleasure.
Whatever conditions obstruct conation in the attainment of

* “Manual of Psychology,” p. 234. “Displeasure’”’ here means the
feeling attitude antithetical to “ pleasure.”

EVOLUTION OF FEELING AND EMOTION 283

its end, are sources of displeasure. This is the widest gene-
ralization which we can frame, from a purely psychological
point of view, as regards the conditions of pleasure and dis-
pleasure respectively.” Here Dr. Stout seems carefully to
avoid the commonly accepted and much advertised conclusion,
that pleasure and pain (to use this more familiar word as
the antithesis of pleasure) are themselves the end of conative
endeavour. And he is so far right that they by no means
constitute the sole or indeed the primary end of all conative
process. Attention is a conative act; but its primary end
is not pleasure, but rather, as Dr. Stout says,* the fuller
presentation of the object. No doubt this brings pleasure ;
but the fuller presentation comes first, and carries the pleasure
with it. Instinctive response to felt stimulus falls within the
conative attitude. In it there is that “inherent tendency
to pass beyond itself and become something different,” which
Dr. Stout assigns to conation as its chief characteristic. But
the end is not pleasure, but simply the instinctive behaviour.
And if we say that the attainment of this end does bring
satisfaction, which is a form of pleasure, Dr. Stout would
probably reply that this is rather a result of the process than
its true end.

Now, in such cases, what we are really dealing with is a
class of organic processes having conscious accompaniments.
No doubt the conscious accompaniments are of importance ;
they certainly cannot be neglected by the psychologist : but
their feeling-tone does not constitute that which makes
instinct run its course. And I have introduced the subject
for present discussion in this way to reinforce what has already
been repeatedly urged in the foregoing pages, that individual
behaviour, in its first intent, is a biological legacy with ends
predetermined through heredity. The inherent tendency to
pass beyond itself and become something different, which for
the old psychology was a heaven-sent impulse, or, as Addison
said, “an immediate impression from the first Mover and the
Divine energy acting in the creatures,” becomes for the new

* Op. cit., p. 65.

284 THE FEELINGS AND EMOTIONS

psychology an organic bequest. But the attainment of ends
thus already predetermined has feeling-tone, both as process
and in its resulting consciousness, and this feeling-tone serves
to modify, through the situation it introduces, future be-
haviour, and thus, in a sense, affords a new end to subsequent
conation.

“ Life,” wrote James Martineau,* “is a cluster of wants
physical, intellectual, affectional, moral, each of which may
have, and all of which may miss, the fitting object. Is the
object withheld or lost? there is pain: is it restored or
gained ? there is pleasure: does it abide or remain constant ?
there is content. The two first are cases of disturbed equi-
librium, and are so far dynamic that they will not rest till
they reach the third, which is their posture of stability and
their true end.” This is an adequate description of the
essential features in conative process. But in genetic pre-
cedence, as in individual development, the physical wants
come first, and, at the outset of behaviour, the satisfaction or
content is not and cannot be foreseen, since it has never yet
entered into experience. To adopt a distinction suggested by
Professor Mackenzie,t the conation is purposive, since we see
that an end is involved, but not purposeful, since there is no
definite consciousness of the end aimed at. But when experi-
ence has introduced feeling-tone into the situation, we may
say that this, in a sense, introduces a new end to subsequent
behaviour.

Mr. Herbert Spencer has said { that pleasure is that which
we seek to bring into consciousness and retain there; pain,
that which we seek to get out of consciousness and keep out.
May we assert, then, that, in the modification of behaviour
due to experience, the pleasure to be gained or the pain to be
avoided is the psychological end? Certainly not without
qualification, unless we be among those who are content to
accept any form of words which gives a general sort of notion

* “ Types of Ethical Theory,” vol. ii, f. 350.
+ “ Manual of Ethics,” p. 85.
+ “Principles of Psychology,” vol. i., pt. ii, ch. ix. § 125,

EVOLUTION OF FEELING AND EMOTION 285

of the kind of thing which we suppose is meant, and which is
probably more or less correct. We want here and now to
get clear ideas, and to express them with some approach to
accuracy. To say that pleasure is the psychological end of
intelligent behaviour is to put the matter too subjectively
and in too abstract a form. Professor Mackenzie has clearly
indicated the ambiguity in the word “pleasure.” “‘ Pleasure,”
he says,* “is sometimes understood to mean agreeable feeling,
or the feeling of satisfaction, and sometimes it is understood
to mean an odject which gives satisfaction. The hearing of
music is sometimes said to be a pleasure, but of course the
hearing of music is not a feeling of satisfaction; it is an
object that gives satisfaction. Generally, it may be observed
that when we speak of ‘pleasures’ in the plural, or rather in
the concrete, we mean objects that give satisfaction ; whereas
when we speak of ‘pleasure’ in the abstract, we more often
mean the feeling of satisfaction which such objects bring with
them.” May we not go a step further, but entirely in the
same direction, and say that pleasure is a constituent part of
the concept self as an object of thought or desire; that its
proper sphere is in the ideational consciousness ; and that, as
we interpret the animal mind, it has no place as such therein ?
The hedonist regards pleasure as the most excellent and dis-
tinctive characteristic of his ideal self and his ideal community.
But animals have not risen or fallen to the level of hedonism.
Pleasure is not for them a motive of conduct, though nice
objects, as such, are attractive, and through them impulse
acquires direction and force. :

If, in animal psychology, we are to use the words pleasure
and pain (as the antithesis of pleasure)—and they seem
more properly to belong to a plane of mental development to
which animals probably have not attained—we may say that
the pleasure or the pain which attaches to any centre of
interest in the situation is that which gives it attractive or
repellent meaning; it furthers conation either towards or,
as Hobbes would say, fromwards. But if we put the matter in

* “Manual of Ethics,” p. 72.

286 THE FEELINGS AND EMOTIONS

this somewhat abstract form, let us keep in view, if it be only
in the background of our thought, the kind of concrete
example which may be adduced in its illustration—the dog
with his attractive bone, the kitten that has raced off at sight
of him, the cock-sparrow with trailing wings hopping after
his mate, the falcon stooping on her quarry, the rabbit diving
into his burrow at sight of the fox, and so forth. If we
have such cases in view, where the centre of the situation
has acquired or is acquiring meaning, a meaning which in
large degree attaches to the external nucleus of the situation
with only the germs of subjective reference, we may, perhaps,
summarize the position by saying that in each case some
pleasure to be gained or some pain to be avoided is the psy-
chological end of conation.

But in each case the conation has also a biological end—
the preservation and conservation of the race. “An animal,”
said Darwin,* “may be led to pursue that course which is most
beneficial to the species by suffering, such as pain, hunger,
thirst, or fear; or by pleasure, as in eating and drinking, and
in the propagation of the species ; or by both combined, as in
the search for food.” The important point here to notice is
that the two ends agree—the psychological end of the attain-
ment of pleasure and the avoidance of pain, and the biological
end of race preservation. Under the joint influence of plea-
sure and pain, the needle of animal life sets towards the pole
of beneficial action.

This consonance of end was in old days ascribed to the
beneficent foresight of the Creator. The modern view, that
it is a product of evolution, does not necessarily ascribe it to
any other ultimate cause. For many still piously hold that
evolution is only a name which we give to the method of
creation. And there is not a fact or generalization in science
by which such a conclusion can be disproved, for the premises
lie outside the field of scientific inquiry. But the consonance
of end is, for science, a remarkable fact, and one worthy of
attentive consideration.

* “ Vife and Letters,” vol. i., p. 310.

EVOLUTION OF FEELING AND EMOTION 287

We have already seen that, if the claim for the inheritance
of acquired characters be, on the evidence, judged unproven,
and if instinct cannot be ascribed to transmitted habit, or re-
garded as a legacy of that which has been ancestrally acquired,
the only scientific explanation of instinctive behaviour is one
which involves the principle of natural selection. But no one
doubts that, in the course of experience, animals acquire modes
of procedure which are beneficial to the race. This is well seen
in the play of animals as interpreted by Professor Groos. Now,
why do animals play ? From the psychological point of view,
because they like it ; from the biological point of view, because
they thus gain practice and preparation for the serious business
of their after-life. But why do they like it? because, under
natural selection, those who did not like it, and therefore did
not play, proved unfit for life’s struggle, and were eliminated.
Suppose that an animal were born with a rooted hereditary
aversion to everything nutritious and an inherited hunger for
anything harmful and unfit for food. What chance would it
stand of survival? Hereditary likes and dislikes determine
the general course of acquired behaviour, just as hereditary
nerve-connections determine the course of instinctive behaviour.
Wherein, then, lies the difference between the two? In the
fact that in the one case the nerve-connections are transmitted
ready-made, while in the other they result from association or
coalescence in the course of individual life. But in both cases
the pursuit and attainment of the beneficial brings satisfaction.

Now, the consonance of end has long been regarded as an
inevitable deduction from the hypothesis of evolution. “ That
pains are correlatives of actions injurious to the organism,”
wrote Mr. Herbert Spencer in his “‘ Principles of Psychology,” *
“while pleasures are the correlatives of actions conducive to its
welfare, is an induction not based on the vital functions only.
It is an inevitable deduction from the hypothesis of evolution,
that races of sentient creatures could have come into existence
under no other conditions. Those races of beings only can

* Vol. i. pt. ii, ch. ix., § 124. I quote from the valuable “ Epitome ”
prepared by Mr. Howard Collins, p. 214.

288 THE FEELINGS AND EMOTIONS

have survived in which, on the average, agreeable or desired
feelings went along with activities conducive to the maintenance
of life, while disagreeable and habitually avoided feelings went
along with activities directly or indirectly destructive of life,
and there must ever have been, other things being equal, the
most numerous and long-continued survivals among races in
which these adjustments of feelings to actions were the best,
tending ever to perfect adjustment.” And he safeguards the
position by adding : “It is frequently taken for granted that
the beneficial actions secured must be actions beneficial to the
individual ; whereas the only necessity is that they shall be
beneficial to the race.”

This aspect of the consonance is now quite familiar ; but
let us carefully note how completely dependent it is on natural
selection. Mr. Herbert Spencer’s testimony is especially valu-
able, since he has always laid much stress on the hereditary
transmission of acquired characters and still holds * “ that the
inheritance of functionally-caused alterations has played a
larger part than Darwin admitted even at the close of his life ;
and that, coming more to the front as evolution has advanced,
it has played the chief part in producing the highest types.”
Now, in these types we certainly find a wide range of consonance
between the psychological and the biological ends of behaviour ;
of which the phenomena of play may again be adduced as an
example. Hence the special value of Mr. Herbert Spencer’s
testimony to the part played by natural selection in establish-
ing the consonance. “Only those races of beings,” he says,
“can have survived in which, on the average, agreeable feelings
went along with activities conducive to life ;” and again, “ The

_ most numerous survivals must ever have been among races in
which these adjustments of feelings to actions were the best.”
The stress is here laid on the survival of those in which the
consonance has obtained ; the elimination of those in which it
was absent: that is to say, on natural selection. And where
else can it be laid? It is not the sort of thing which could be
acquired. Suppose that, as we suggested above, an animal were

* “Principles of Biology,” revised and enlarged edit. (1898), p. 560.

EVOLUTION OF FEELING AND EMOTION 289

born with a rooted hereditary aversion to everything nutritious
and an inherited hunger for anything harmful and unfit for
food. Under what conceivable conditions could such an animal
acquire a complete change of its affective nature? Animals
like things or they do not like them ; only to a very limited
extent, if at all, under natural conditions, can they learn to
like them. We, indeed, can in some degree learn to take
pleasure in that which at first, and by nature, is distasteful ;
but we do so by some external constraint, or from some motive
of ideational origin. We put pressure upon ourselves, or have
_ pressure put upon us, repeatedly to perform some irksome task ;
we fall into routine and custom ; and the performance becomes
so far second nature that its discontinuance produces an un-
comfortable sense of something lacking in the daily round.
Perhaps domestic animals learn to like the good offices we force
them to perform for us. But here we have the element of
external constraint, which is wholly, or almost wholly, absent
under natural conditions. And there is no evidence that such
acquired likings are inherited. That, however, is another
question. Our present point is that, under nature, the condi-
tions of such acquisition are lacking ; so that, there being no
acquisition, there is, in this case, nothing acquired to be
transmitted.

But, so far as behaviour is concerned, “ functionally
caused alterations” are those due to the exercise of intelli-
gence, by which the behaviour acquires direction and character
in reference to the meaning introduced into the situations.
See, then, the position to which we are logically driven. The
acquisition of that which has beneficial value in behaviour
depends on a consonance between psychological and biological
end. But this consonance is dependent on survival, and,
apart from special creation, or some kindred hypothesis such
as Leibnitzian harmony, can be due to nothing else. Even if
we grant, therefore, that the effects of acquisition are inherited,
the conditions of beneficial acquisition are dependent on
natural selection. And thus the inheritance of acquired

characters, which is so often urged as a principle of evolution
U

290 THE FEELINGS AND EMOTIONS

independent of natural selection, is, so far as intelligent
behaviour is concerned, indirectly, if not directly, due to this
very natural selection of which it is said to be independent.
Surely, under these circumstances, the hypothesis in question
may be said to be not only unproven, but altogether un-
necessary.

And what is true of those diverse feelings which we group
under the concepts pleasure and pain respectively, is true also
of those more complex dispositions which we call emotional—
using this term in a broad and comprehensive sense. We say
that in their primary manifestations they are instinctive ; and
they certainly seem to accompany organic behaviour due to
co-ordinated reflex actions. But the emotion, as instinctive, is
a matter only of its first occurrence. In the course of ex-
perience it enters into conscious situations, the centres of
interest in which have acquired meaning.

Take a particular case.* Your dog is dozing on the lawn
in the sunshine. Suddenly he raises his head, pricks his ears,
scents the air, looks fixedly at a gap in the hedge, and utters a
low growl. Place your hand on his shoulder, and you will
find that his muscles are all a-tremble ; on his ribs, and you
will feel how strongly his heart is beating. Soon the growing
excitement leads to vigorous action, and he darts through the
gap. You follow him across the lawn, look over the hedge,
and see him facing his old enemy, the butcher’s cur. They
are moving slowly past each other, head down, teeth bared,
back roughened. You whistle softly. Such a call would
generally bring him bounding to your feet; but now it is
apparently unheard, at any rate unheeded. The two dogs
have a short scuffle, and the cur slinks off. Your dog races
after him ; and he flees, yelping. The situation is over. Spot
returns, wagging his short tail, jumps up at you playfully,
and then lies down again on the grass. But now and then,
for ten minutes or so, he raises his head and growls softly.

Let us briefly analyze the dog’s condition and actions,
reading into them, conjecturally, the accompaniments in

* From “ Animal Life and Intelligence,” p. 382.

EVOLUTION OF FEELING AND EMOTION 291

consciousness. As he lies on the lawn, he receives a sense-
stimulus, auditory or olfactory. It has already acquired
meaning, from many a tussle with the butcher’s cur. It
has organic effects, and it generates a conscious situation
which has acquired complexity through coalescence. As the
result of this situation the head is raised, the ears pricked, and
soon, The dog is on the alert. His attention is aroused.
The muscles of neck, eyes, ears, are brought into play in such
away as to bring the senses to bear on the exciting object.
He probably sees the cur through the gap in the hedge. The
muscles of the frame are innervated so as to be ina state of
preparation to act rapidly and forcibly. At the same time the
vaso-motor system is disturbed, the heart-beat is quickened,
respiration is altered ; there is probably hardly an organ in
the body which remains unaffected. Then the dog rushes
through the hedge, and stands with bared teeth before his
antagonist. A whole set of appropriate muscles are now
strongly innervated. There is, perhaps, a double innervation,
stimulating to activity and yet restraining from action. He
bares his teeth and growls deeply. Attention is so concen-
trated that he heeds not, perhaps does not hear, his master’s
whistle. He is keenly on the alert. The blood-system, respi-
ratory organs, and all his inner machinery are still pulsating
with nervous thrills ; his backis up. Then he sees his chance,
and flies at his opponent. Much that he has learnt in play,
and all that he has learnt in earnest, comes to his aid in the
short angry scuffle. And what we call his emotion of anger
spurs him on to the fight; the cowardly dog in which this is
lacking or is replaced by fear is spurred to flight. Hach may
contribute to self-preservation, but in different ways.

Now, we shall not attempt to determine how the distinc-
tively emotional elements arise. Some think they arise by a
sort of irradiating nervous diffusion in the nerve-centres as
a direct result of the originating stimulus. Mr. Rutgers
Marshall regards them as due to the motor activities in fight
or flight ; Professor William James contends that they have
their source in the visceral affections of heart, lungs, glands,

292 THE FEELINGS AND EMOTIONS

and so forth ; Professor Lange attributes them to vaso-motor
effects. The problem is a difficult one, and hard to determine
by experiment ; for we have to deal with a matter of primary
genesis, of how they are at the outset introduced into the
conscious situation. Experiments on animals which have
already gained emotional experience cannot decide the ques-
tion of genesis. Professor Sherrington, for example, has
shown * that, after severance of the spinal cord in the lower
region of the neck, and of the vagus nerves, by which “a
huge field of vascular, visceral, cutaneous, and motor re-
action” were “deprived of all connection with the nervous
centre necessary to conscious response,” “the emotional states
of anger, delight at being caressed, fear and disgust were
developed with, as far as could be seen, unlessened strength.”
But the avenues of connection were closed after the motor and
visceral effects had played their parts in the genesis of the
emotion on the hypothesis that the emotion is thus generated.
Although new presentative data of this type were thus excluded,
their re-presentative after-effects in the situation were not ex-
cluded. It is, moreover, an essential part of Professor James’s
doctrine, as I provisionally accept it, that the ‘ expression ” and
the visceral and vascular efforts are independent results of stimu-
lation in certain ways, and that these independent results are
conjoined through natural selection. Suppose we sever the
connection through which the one takes effect, there is no
reason to expect that the manifestation of the other would
cease. Professor Sherrington cut off the channels of com-
munication with the visceral and vascular apparatus: if the
channels of expression remained open there is no reason why
such expression should cease.

We need not, however, for our present purpose, attempt to
ascertain how the distinctively emotional characteristics arise.
It is sufficient that they are presumably present in the situa-
tion. Now, as Dr. Stout well points out,t the emotions

* “Experiments on the Value of Vascular and Visceral Factors for

the Genesis of Emotion,” Proc. Roy. Soc., vol. Ixvi., pp. 390-403 (1900),
+ “Manual of Psychology,” p. 288.

EVOLUTION OF FEELING AND EMOTION 293

generally presuppose the existence of certain specific ten-
dencies. ‘The anger produced in a dog by taking away its
bone presupposes the specific appetite for food. The anger
produced in it by interfering with its young presupposes the
specific tendency to guard and tend its offspring. So the
presence of a rival who interferes with its wooing causes
anger because of the pre-existence of the sexual impulse.” In
general, we may say that emotional states are, under natural
conditions, closely associated with behaviour of biological
value—with tendencies which are beneficial in self-preserva-
tion or race-preservation—with actions that promote survival,
and especially with the behaviour which clusters round the
pairing and parental instincts. The value of the emotions in
animals is that they are an indirect means of furthering sur-
vival. But how has the close association between emotional
condition and the biological end it furthers been established?
Again, we must say that under natural conditions it is not the
sort of thing which could be acquired. And again we must
urge that natural selection through survival is, apart from
some theory of pre-established harmony, the only hypothesis
in the field on which the close association can be explained.
There is one more point to which attention may be drawn.
If there be one thing, and there certainly are not many, on
which all writers on the emotions are agreed, it is as to their
vagueness. They do not readily submit to definition, and
cannot be described in a sentence. ‘This is not due to any
indefiniteness of biological end, nor to much indefiniteness in
the mode of “expression ;” it is due, rather, to an inherent
dimness and haziness of psychological outline. We seem
unable to focus them and get a clear-cut result. This is, no
doubt, in part due to the complexity of emotional states. But,
may it not be largely due to the fact that there is no necessity
for definiteness? They fulfil their purpose just as well if they
are vague. It is quite necessary for the dog to have a clear-
cut impression of his antagonist ; and, on the cognitive side
of consciousness, meaning must be in some degree definite
to be of real value. But, so long as the emotion raises the

204 THE FEELINGS AND EMOTIONS

temperature, so to speak, to the boiling-point of vigorous
action, ib matters little what the psychological source of heat
may be. If this be so, we should expect an emotional vague-
ness, since natural selection puts no premium upon emotional
definiteness. And from this it follows, as a corollary, that,
whereas we may infer that an animal’s perceptual products are
probably closely similar to our own, since sight, touch, hearing,
smell, and taste are of value in so far as they convey definite
meaning, in interpreting their feelings and emotions we
have less secure grounds of inference, since all that is requisite
is that there should be a sufficiently high emotional tempera-
ture to afford the conditions for definite and vigorous action.
In conclusion, then, we may say that the primary purpose _
of the evolution of fecline and emotion is to promote beneficial
behaviour, | and. that the observed consonance of the psycho-
logical “end ¢ of attaining satisfaction, and the biological end_ot_
securing survival, seems to be due to natural selection—is,
indeed, scarcely explicable on any other naturalistic hypothesis.
A word of warning may be added. We have repeatedly
spoken of biological and psychological ends. By this we mean
what seems to the observer, as an interpreter of natural pro-
cesses, the purpose and object of their existence. But the
word “end” is often used in such a way as to imply foresight
and contrivance on the part of a rational being. We have
not used it in this sense. Whether the whole of nature, in-
cluding animal behaviour, is driven onwards to definite ends
by an underlying Cause, is a metaphysical question. It is not
one on which science has any right to express an opinion one
way or the other. Science deals with the phenomena ; the
causes of their being lie outside her province.

CHAPTER VII
THE EVOLUTION OF ANIMAL BEHAVIOUR

I.—TueE PuysriotogicaL ASPECT

AT the outset of our inquiry, we used the word “ behaviour ”
in a wide and comprehensive sense. Thus broadly used, I
said, the term in all cases indicates and draws attention to the
reaction of that which we speak of as behaving in response
to certain surrounding forces or circumstances which evoke
the behaviour. The behaviour of living cells is dependent on
changes in their environment; that of deciduons trees, as
they put forth their leaves in the spring or shed them
in the autumn, is related to the change of the season; in-
stinctive, intelligent, and emotional behaviour are called forth
in response to those circumstances which exercise a constrain-
ing influence at the moment of action. Used in this com-
prehensive sense, the term “behaviour” neither implies nor
excludes the presence of consciousness. We know from our
own experience, however, that consciousness does in some
cases accompany behaviour, and we infer that in many other
cases it may be present. But we need a criterion of its
presence to guide our inferences, and this criterion we found
in the ability of living beings to profit by experience. In
Dr. Stout’s phraseology, if a thing seems to acquire meaning
for such a being, and the behaviour is guided in accordance
with such acquired meaning, we infer the presence of con-
sciousness as supplying conditions effective in determining
its course. Still this does not exclude, nay, rather it pre-
supposes, the presence of sentience at a lower stage of evolution,
295

296 THE EVOLUTION OF ANIMAL BEHAVIOUR

a sentience which is as yet ineffective since the process of
conscious coalescence has not begun, or has not been carried
far cnough.

In foregoing chapters we have constantly held the problems
of evolution in view, and in special sections directed attention
to them. But the subject is so central to modern thought and
discussion, that some further consideration of certain aspects
of the evolutionary process and products will fitly serve to
bring our inquiry to a conclusion.

We must accept, asa datum from the physiological point
of view, the fact that protoplasm does respond to stimuli,—that
it possesses the fundamental property of irritability. It is a
substance that is in a state of unstable equilibrium. Its ten-
dency to pass to a condition of more stable equilibrium is that
in and through which organic behaviour in its very simplest
expression is possible. And this, with progressive complication,
runs through the whole gamut of animal behaviour, and
eventually passes over into the sphere of consciousness. ‘‘ The
tendency to equilibrium,” writes Dr. Stout,* ‘is the physio-
logical correlate of what on the psychical side we call conation,
—the striving aspect of consciousness.” But, protoplasm at the
outset—or as near the outset as we can get—is, in technical
phrase, differentially responsive. The nature of the stimulus
and the nature of the conditions decide what the nature of
the response shall be. And even in that jelly-like speck of
living matter, the Amada, the responses conspire to a biological
end. If they did not so conspire, we should not have the
phenomena of life. The mere act of living, building up from
food-stuff and oxygen an unstable substance which “ ex-
plodes” and contracts under stimulation, implies that the
processes which thus conspire are related in such a manner as
to fulfil and secure their end. In higher unicellular animals,
such as the Paramecium, the relations are less simple ; but in
them the continuance of that sum of organic behaviour which
we call life, is secured only on the condition that these less
simple relations are duly preserved, and that the vital processes

* “ Manual of Psychology,” p. 132.

“

THE PHYSIOLOGICAL ASPECT 297

conspire with sufficient unity of biological purpose. And
when we pass to the higher creatures in which many cells
unite to form one animal, the very word “unite” indicates
that the vital processes of all must conspire with sufficient
unity of biological purpose to insure the continued life of the
whole.

Now, in all the higher and more active animals a nervous
system is developed, which has for its purpose and end the
preservation and furtherance of unity amid circumstances of
progressively increasing diversity. And in the course of its
evolution an added means of preserving and furthering the
essential unity is provided in consciousness, which, through the
coalescence of scattered units of sentience, leads behaviour to
acquire a new and higher unanimity of purpose. Thus a
mental evolution is engrafted on the organic evolution which
precedes it. But every step in this mental evolution presup-
poses a step in organic evolution. And such is the complexity
of structure and process in all the higher animals that much
of the business of behaviour is relegated to quasi-independent
nervous centres, which perform this business automatically, and
will continue to perform it, with much subsidiary unity of end,
when they are left to themselves and all connection with the
supremely unifying sensorium has been severed.

- Before proceeding to give some examples of this fact, and
to indicate its bearing on our interpretation of behaviour, it
may be well to state distinctly that no attempt is or will be
here made to trace in detail the course of the evolution of
animal behaviour through the ascending grades of life, nor,
indeed, to prove that there has been any such evolution.
Evolution by natural genesis is here assumed as the only
hypothesis with which science has any concern. If it be false,
then have the labours of workers and thinkers, since Darwin
and Mr. Herbert Spencer worked and thought, been vain.
Special creation is not a scientific hypothesis, but a reference
of biological and mental phenomena to an ultimate cause,
which lies beyond and altogether apart from the scope of
scientific inquiry. The fundamental assumption of the man

298 THE EVOLUTION OF ANIMAL BEHAVIOUR

of science is, that any natural event he may select for detailed
study has natural conditions and antecedents. And ib is only
in such detailed study—taking this or that particular occur-
~ rence and endeavouring to ascertain what were its related
antecedents—that advance in the evolutionary interpretation
of nature can be secured.

Such advance has been secured by the labours of those
physiologists who have established by careful experiment the
quasi-independent action of subsidiary nerve-centres as con-
stituents of the nervous system as a whole. In such animals
as the crayfish and the lobster the central nervous system
consists of a chain of “ ganglia,” or nerve-knots, which are
connected together by nerve-strands. If these strands be cut
between the thorax, which carries the walking limbs, and the
abdomen or hinder portion of the body, the nerve-connection
between these parts is severed. If the forepart be irritated
through its sense-organs, the limbs of that part will respond ;
but, whereas an unmutilated crayfish, subjected to such irrita-
tion, would give a vigorous flap of the tail, this does not take
place in the crippled animal.* Still, if the abdomen be
irritated, it will respond by a strong and swift contraction.
The two portions of the body are each capable of acting
independently with well co-ordinated movements, but no longer
of working together with unity of purpose. In the hinder
portion the abdominal limbs, or swimmerets, all swing back-
wards and forwards simultaneously with rhythmic strokes ;
they act in concert. Sever now the connections between their
ganglia, and each pair of limbs will continue to swing
rhythmically but not with concerted rhythm. We have isolated
a number of quasi-independent centres, and rendered them
really independent. Hach is concerned with its own proper co-
ordination, but can no longer combine with others in a wider
co-ordination. Mr. Hydef has shown that in the king-crab,

* See Huxley’s book on ‘The Crayfish,” in the International Science
Series, p. 108.

+ Journal of Morphology, vol. ix. Quoted by Professor C. S.
Sherrington in The Marshall Hall Address, “On the Spinal Animal ”
(reprinted from Medico-Chirurgical Transactions, vol. 82), p. 4.

THE PHYSIOLOGICAL ASPECT 299

Limulus, when the nerve-chain is severed just in front of the
abdominal region, the rhythmic respiratory movements of the
abdominal segments still proceed regularly and co-ordinately.
Even when only a fraction of the nerve-cord, separated by
severances in front and behind, is left, corresponding with a
single abdominal segment, the rhythmic action of that segment
continues ; but it is no longer synchronous with that of
adjacent segments similarly isolated.

It will probably not be contended that the co-ordinated
rhythm of the isolated seement in crayfish or king-crab is any-
thing but a bit of organic and physiological behaviour.
Whether it be accompanied by consciousness—a bit of con-
sciousness isolated from other bits—we do not know; but we
have no grounds for supposing that the rhythmic behaviour is
guided by consciousness. And when, as Dr. Carpenter pointed
out half a century ago, a water-beetle, from which the ‘“ brain ”
has been removed, swims forwards if placed in water, we must
surely regard the co-ordinated progression as organic behaviour,
whatever view we may hold with regard to a consciousness
which in such a case is in a very literal sense a divided
consciousness.

In these invertebrates the central nervous system is obviously
segmented—one can distinguish the ganglia and their con-
necting nerve-strands. In the vertebrate the brain and spinal
cord form a continuous mass of nerve-tissue without obvious
segmentation. But the pairs of spinal nerves, each nerve with
its afferent and efferent ‘‘root,” indicate a really segmented
condition, though in the cord itself the segments so run
together and overlap that they cease to be externally obvious.
And there is a certain, though limited, amount of overlap in
the distribution of these segmental nerves. Still, well co-
ordinated responses occur when comparatively short portions
of the spinal cord are isolated by severance from the rest. In
the male frog, especially during the breeding season, a clasping
reflex is produced by stimulating the dark swollen pads on the
inner side of the hand, and this, as Goltz has shown, is
exhibited when all the central nervous system has been destroyed

300 THE EVOLUTION OF ANIMAL BEHAVIOUR

save the segments to which the nerves for the arms proceed.
“ Similarly,” writes Dr. Sherrington,* “in the cat and monkey,
the reflex wagging of the tail persists when behind the spinal
transection only the sacral region of the cord is left intact.”
When the spinal cord of the dog is severed, so as to isolate
that portion which is concerned with movements of the hinder
part of the body, pressure on the pad of one hind-foot usually
produces, not only a lifting of that leg, but also an extension
of its fellow—that is to say, a co-ordinated response of the two
limbs. But in the case of the vertebrates, more than in that
of the invertebrates, the co-ordinated response of an isolated
part of the central nervous system seems to lack the furtherance
of its action, which normally comes from the higher centres
from which it has been severed. ‘The spinal reflexes signifi-
cant of progression seem,” says Dr. Sherrington,t “to con-
tribute chiefly towards preparatory posture in readiness for the
onset of action executed by the musculature under the driving
of higher centres. Thus the well-known reflex spinal posture
of the frog is flexion of the hind limbs, the extensors of the
joints being taut and ready for the jump. ‘The spinal reflexes,
which in their results approximate most closely to the normal
reactions of the unmutilated individual, are those connected
with the pelvic and abdominal viscera,” many of which “are
executed as spinal reflexes in a manner presenting little or no
physiological defect from the normal. And if the bulb ”—the
continuation of the spinal cord within the skull to form the
basal portion of the brain—“ be included with the spinal cord,
and these together, including their nerves, be isolated from the
rest of the nervous system, the animal as regards its visceral
life, including that of the heart and lungs, is practically intact.”

Huxley graphically describes the actions of a frog from
which the cerebral hemispheres have been removed. “If that
operation,” he says,{ “‘is performed quickly and skilfully, the

* «The Spinal Animal,” p. 5.

t Op. ctt., p. 28.

$ “Collected Essays,” vol. i, essay on ‘ Animal Automatisin,” p.
224,

THE PHYSIOLOGICAL ASPECT 301

frog may be kept in a state of full bodily vigour for months, or
it may be for years ; but it will sit unmoved. It sees nothing ;
it hears nothing. It will starve sooner than feed itself,
although food put into its mouth is swallowed. On irritation,
it jumps or walks; if thrown into water it swims. If it be
put on the hand it sits there, crouched, perfectly quiet, and
would sit there for ever. If the hand be inclined very gently
and slowly, so that the frog would naturally tend to slip off,
the creature’s fore paws are shifted on to the edge of the hand,
until he can just prevent himself from falling. If the turning
of the hand be slowly continued, he mounts up with great care
and deliberation, putting first one leg forward and then
another, until he balances himself with perfect precision on the
edge ; and if the turning of the hand is continued, he goes
through the needful set of muscular operations, until he comes
to be seated in security on the back of the hand. The doing
of all this requires a delicacy of co-ordination and a precision of
adjustment of the muscular apparatus of the body which are
only comparable to those of a rope-dancer.”

Now, why have we entered into these details ? To rein-
force, from a somewhat different point of view, that which has
again and again been urged in the preceding sections of this
inquiry, that much of animal behaviour is an organic legacy.
A going mechanism of great delicacy, with ready-made co-
ordinations, the products of biological evolution, affords to
consciousness a vast body of its primary data. As Dr, Sher-
rington himself says,* “ co-ordination is abundantly shown to
result from the independent power of the spinal arcs, altogether
apart from the influence of the great cranial sense-organs, and
of the cerebral arcs superposed upon them. These senses and
the brain find the elementary co-ordination of the skeletal
musculature an achievement already provided and to hand in
the spinal cord. And no doubt the product of the instrument
is, with the instrument itself, given over to their use in the
reactions they elicit from the spinal musculature.” We have
seen how instinctive behaviour, in those animals in which it is

* Op. cit., pp. 20, 21.

302 THE EVOLUTION OF ANIMAL BEHAVIOUR

best. studied, affords in hereditary biological outline, a sketch
which subsequent acquisition, under experience, serves only to
elaborate by the filling in of details and of the more delicate
shading in behaviour. But in the higher animals, in which a
period of youth is a time for the acquisition of experience—for
experimentation and practice,—it might seem that the inherited
biological legacy was of less importance. The “spinal animal,”
as Dr. Sherrington calls it,—that is, the animal in which the
spinal centres are isolated from the cerebral centres—goes far
to disprove any such view. In them the cerebral senses and
the brain find elementary co-ordination of the bodily move-
ments an achievement already provided and to hand in the
spinal cord. But when different animals are compared—frog,
bird, rabbit, dog, and monkey—the permanent effects of sever-
ance of brain connection, the effects which remain when the
temporary period of disturbing “shock” is over, are more
marked in the higher than the lower types. And concerning
this, Dr. Sherrington says,* “The deeper depression of reaction
into which the higher animal, as contrasted with the lower,
sinks when made spinal, appears to me significant of this, that
in the higher types, more than in the lower, the great cerebral
senses actuate the motor organs, and impel the motions of the -
individual.”

Whether in the animal in which all direct connection be-
tween the anterior and posterior portions of the central nervous
system has been severed there is a double consciousness—a
cerebral consciousness and a spinal consciousness—we are not
in a position accurately to determine. If the generally accepted
opinion, that the higher brain-centres constitute the sensorium
or seat of consciousness, be correct, we must suppose that a
maimed consciousness, with many avenues of experience closed,
is retained in the anterior moiety, while the posterior is rele-
gated to a condition of mere sentience at best. In any case all
relation between the two is prevented. The two—if two there
be—are rendered quite independent through severance of the
cord in the region of the neck. But there is a point of view

* Op. cit., p. 29.

THE PHYSIOLOGICAL ASPECT 303

indicated by Dr. Sherrington which is full of suggestion and
interest.

“Tt is significant in the evolution of animal form,” he
says,* ‘‘that the organ that exhibits most uninterrupted and
harmonious increase in development, as studied successively in
passing from lowest to highest, is the brain. And it is signifi-
cant that in the nervous system—segmental system as it is—
the brain. is developed, not in those segments whose sense
organs are ordinary cutaneous (tactual, etc.), muscular and
visceral, but in the segments connected with the visual, olfac-
tory, and auditory sense-organs ; in other words, the brain is
developed in the head. The head is, so to say, the individual ;
it has the mouth, it takes the food, including air and water,
and it has the main sense organs providing data for both space
and time. To this the body, an elongated motor organ with a
share of the viscera and the skin, is appended primarily as
a machine for locomotion. This latter must of necessity lie at
the behest of the great sense organs of the head.”

Now let us try and picture to ourselves a spinal animal, or
one which retains only the lowest portion of the brain, the
part known as the bulb or medulla oblongata; let us assume
that it is conscious and capable of acquiring experience through
the association and coalescence of the data afforded by the
senses that remain to it; and let us try to imagine the con-
scious situations which would arise, and their value in the
guidance of behaviour. The senses that remain are touch and
the temperature sense, the motor sense affording data from the
muscles, joints, and tendons, and those which supply certain
visceral sensations. There is not one of much, if any, guiding
value left. There is not one of what we may term anticipatory
use. There is not one which could serve to infuse anything
like definite meaning into the situation. For, after all,
meaning is expectation. There is an element of anticipation
in all those senses which are of any real guiding value in
the conscious situation. Sight, hearing, and, especially for
some animals, smell,—these are the senses which forewarn of

* Op. cit., p. 18.

304 THE EVOLUTION OF ANIMAL BEHAVIOUR

something which may follow; of other sensations with which in
the course of experience they have coalesced. And they are all
cut off from the supposed spinal animal. A light touch might
in some cases forewarn of the shock or severe pressure, which
would perhaps follow. But the shocks would so often come
suddenly that it is questionable whether the warning would be
of much avail. Still, touch is a warning and cognitive sense,
and through it the environment would acquire -a limited
amount of meaning.

Now, the biological value of coalescent association lies in
this very element of warning. The anticipatory senses, sight,
hearing, smell, are in their several degrees the “ projective ”
senses, the senses which carry with them the quantity of “ out-
ness.” And their “projective” character is the necessary
psychological expression of their distinctive biological end.
They must be projective, must carry with them “ outness,” if
they are to convey what we, following Dr. Stout, have so often
spoken of as meaning. But if the biological value of co-
alescent association lies in the expectation it renders possible ;
and if, in the spinal animal, there are no senses left save touch,
which could receive from the environment preparatory warning
of what is coming ; it would seem exceedingly improbable that
it should develop quasi-independent conscious situations of its
own. In the unmutilated animal, at any rate, tactual ex-
perience would most probably coalesce with that derived from
the senses which more distinctively take the lead in the
acquisition of meaning. And we may therefore, on these
grounds, as well as on others, acquiesce in the current view
that the quasi-independent functioning of the spinal cord and
its constituent segments is, at best, lit up with those flashes of
mere sentience of which Sir Michael Foster speaks in the pas-
sage we quoted in an earlier section.*

If from the consideration of the isolated spinal animal
we turn for a moment to that of the isolated cerebral animal,
we find it in a very different position. It is possessed of the
warning senses—those which from several points of view are

* Vide supra, p. 33.

THE BIOLOGICAL ASPECT 305

>

they have only a very limited company to conduct into action.
The company is there, on the further side of the severance,
but they have lost touch with it. They know not what it is
doing, and have therefore neither the data nor the executive
power to guide its manceuvres in the field of behaviour.
They can form maimed coalescent situations, but they are
as impotent as a mere theorist devoid of all power of practical
application. We need not, however, follow the theme further.
We need only add that, could we isolate tracts of nervous
tissue in the lower brain-centres of such a cerebral animal,
‘we should find that subsidiary co-ordinations would belong,
as a physiological heritage, to these isolated fragments.

The conclusions we may draw, then, with regard to the
evolution of behaviour, as viewed in its physiological aspect,
are that it is, in its simplest expression and in its most
complex, conditioned by sufficient unity of purpose to meet
the biological end of survival; that the complex unity of
purpose may be analyzed into a multiplicity of subsidiary
processes each with its subsidiary unity of purpose; and that
the psychological coalescence which gives unity to experience
under the guidance of the leading senses, is paralleled in
a physiological coalescence within the nexus of the nervous
system.

the leading senses—but they are leaders without a following ;

IJ.—Tux Brotogican ASPECT

The biological aspect of behaviour—its relation to biological
ends—has so often come under our consideration in the fore-
going chapters that little need be added in this section : and
that little may be most appropriately devoted, first to the
question whether consciousness does influence behaviour ; and
secondly, this being accepted, to the importance of the réle
that is played by the development of conscious situations in
securing, in the higher animals, the biological end of racial
preservation.

That this end is secured without the aid of consciousness

x

306 THE EVOLUTION OF ANIMAL BEHAVIOUR

in the case of many organic species, in all those, for ex-
ample, which we classify as plants, must not be taken as pre-
sumptive evidence that in other species, for instance in the
multitudinous host of insects, the development of conscious
situations is of no biological value. The fact that chlorophyll
is not developed in any mammal does not show that the
possession of this substance is of no service to the higher plants.
It would not be worth while to give expression to this very
obvious truth, were it not that critics of natural selection
persistently argue that because one species gets on perfectly
well without this or that particular character it can have
played no part in securing the survival of another species.
When I described, at a meeting of naturalists, how well young
chicks could swim, such a critic drew me aside after the
meeting, and expressed his surprise that this did not convince
me that the webbed foot of the duck could not logically
be attributed to natural selection. This is an extreme case,
and one obviously taken on peculiarly weak grounds. But
even Huxley urged that, because a frog, from which the
cerebral hemispheres have been removed, performs many
co-ordinated actions without conscious guidance, consciousness
is, throughout nature, merely an accompaniment of certain
molecular changes in the brain. “Such a frog,” he says,*
“walks, hops, swims, and goes through his gymnastic per-
formances quite as well without consciousness, and consequently
without volition, as with it; and if a frog, in his natural
state, possesses anything corresponding with what we call
volition, there is no reason to think that it is anything but
a concomitant of the molecular changes in the brain which
form part of the series involved in the production of motion.
“The consciousness of brutes,” he continues, ‘ would
appear to be related to the mechanism of their body simply
as a collateral product of its working, and to be as completely
without any power of modifying that working as the steam-
whistle which accompanies the work of a locomotive engine is
without influence on its machinery. Their volition, if they
* “Collected Essays,” vol. i., p. 240.

THE BIOLOGICAL ASPECT 307

have any, is an emotion indicative of physical changes, not
a cause of such changes. It does not enter into the chain of
causation of their actions at all.”

If the literal truth of this contention—the logical sound-
ness of this conclusion—be admitted, it seems absurd to speak
of the biological value of consciousness in behaviour or to
discuss the importance of the réle that is played by the
development of conscious situations in securing the biological
end of racial preservation.

Now, consciousness is regarded by an influential school of
thinkers as a sort of deus ex machina, which, sitting enthroned,
and crowned with a capital letter as Will, directs, like a being
from another sphere, the doings of the body. It was against
the doctrines of this school that Huxley took up arms. They
do not concern us here. The will, or volition, as an under-
lying cause, stands outside the pale of scientific inquiry. It
belongs to the wide realm of metaphysics; its plea must be
heard in another court. In this part of his contention Huxley
was, we believe, unquestionably right from the scientific stand-
point. Neither will, nor impulse, nor instinct, nor conscious-
ness itself, should be introduced into any scientific description
or explanation of phenomena as a cause of their existence or
being, for as such it does not enter into the sequence of
events; it is that which metaphysics claims as their raison d’étre
—that which gives them being. Science in this matter should
be frankly agnostic—neither affirming nor denying aught.
This, of course, is not equivalent to saying that the agnostic
position is the true end of human reason. That would only
be so on the assumption that the problems of science are the
only problems with which that reason can deal. To exclude
metaphysics from science is not to exclude it from human
thought. As a matter of fact such exclusion is neither possible
nor reasonable. But to clearly distinguish the problems of
science from those of metaphysics is absolutely necessary, if
we are to prevent hopeless confusion of issues.

In contending, however, against the introduction of meta-
physical doctrines into the region of scientific explanation,

308 THE EVOLUTION OF ANIMAL BEHAVIOUR

Huxley seems to have been carried too far by the force of his own
attack. So loug as he held to the position that every conscious
state has, as its concomitant, a molecular change in the brain,
he had all the forces of evolution on his side. But when he
said that consciousness is merely the steam-whistle of life’s
locomotive, or merely answers to the sound which the animal-
bell gives out when it is struck, he takes up another position
of far less strategical strength. For whereas the frog from
which the physical centres of consciousness have been removed
sits crouched and motionless, and “will starve sooner than
feed itself, although food put into its mouth is swallowed ; ”
the frog in which conscious situations can take form in
unmutilated cerebral hemispheres behaves in a very different
manner. It is nothing less than pure assumption to say
that the consciousness, which is admitted to be present, has
practically no effect whatever upon the behaviour. And we
must ask any evolutionist who accepts this conclusion, how
he accounts on evolutionary grounds for the existence of a
useless adjunct to neural processes.

“Tt is,” says Huxley,* ‘ experimentally demonstrable—any
one who cares to run a pin into himself may perform a
sufficient demonstration of the fact—that a mode of motion
of the nervous system is the immediate antecedent of a state
of consciousness. We have as much reason for regarding the
mode of motion as the cause of the state of consciousness, as
we have for regarding any event as the cause of another.
How the one phenomenon causes the other we know as much,
or as little, as in any other case of causation ; but we have
as much right to believe that the sensation is an effect of the
molecular change, as we have to believe that motion is an
effect of impact; and there is as much propriety in saying
that the brain evolves sensation, as there is in saying that
an iron rod, when hammered, evolves heat.” But if we speak
of the related antecedent as the cause, it is not obvious why
we should not describe the desire to demonstrate the supposed
fact as the cause of running in the pin. Weszem to have just

* Op. cit, pp. 238, 239.

THE BIOLOGICAL ASPECT 309

as much reason for calling this antecedent state of consciousness
the cause of certain movements and behaviour, as of calling a
mode of motion in the brain the cause of a further state of
consciousness. It is true that we have not the least idea how
the desire can cause the act; but Huxley practically admits
that we have no idea how molecular change can be the cause
of consciousness. In the one case we are no worse off than
we are in the other. Neither position is logically defensible ;
since each assumes that physical events and states of con-
sciousness can constitute links in the same causal chain.

The philosophical hypothesis known as monism regards
the molecular change, not as the antecedent of a conscious
state, but as its concomitant. That which from a physical
and physiological point of view is a complex molecular dis-
turbance is, at the same time, from a psychological point of
view, a state of consciousness. The two are different aspects
of one natural occurrence. Why such an occurrence should
have two so different aspects we have not the faintest idea ;
but here we are not one whit worse off than we were before.
The hypothesis does, however, help us to get over our difficulty.
An essential feature of Huxley’s contention is that the physical
and physiological chain of causation is complete in itself,
which may be granted ; and further, that if consciousness does
arise it is merely an adjunct without influence on the sequence
of events—what is influential is the molecular disturbance, not
the consciousness which accompanies it. But according to
monism the state of consciousness actually is that very same
something which the physiologist calls, in the language of
physics, a molecular disturbance. And in saying that conscious-
ness influences behaviour one who accepts this hypothesis
is merely avoiding a cumbrous form of circumlocution. He
puts it in this way instead of saying that the nerve-changes in
the cerebral hemispheres, or elsewhere, which from a psycho-
logical point of view are a conscious situation, influence and
determine the course of behaviour. But from this point of
view it is absurd to say that the consciousness is merely an
adjunct—absurd to say that were there no conscious situation

310 THE EVOLUTION OF ANIMAL BEHAVIOUR

the neural situation would remain unchanged. They are the
very same thing from different points of view; and to say
there is no influential conscious situation is simply equivalent
to saying that there is not this determining neural situation.

However we explain the fact, there are few who hesitate
to accept it for the purposes of scientific explanation. The
conscious situation, having no doubt for the physiologist a
neural aspect if he could only get at it as a whole, does
practically determine the behaviour of the animal which has
gained the requisite experience. If we accept the fact, we
may pass on to its importance in securing the biological end
of race preservation.

It is a commonplace of evolutionary doctrine that, other
things being equal, those races will survive, in the constituent
members of which intelligent behaviour enables them to deal
most effectually with an environment of increasing complexity.
And it is a matter of familiar observation that such behaviour
is closely connected with delicacy and refinement of develop-
ment in those senses which take the lead in cognitional process,
and with rapidity and precision in the motor co-ordination
through which prompt and skilful advantage is taken of the
situation which has, through experience, acquired meaning.

But though the importance of intelligent adjustment to
the circumstances of life is widely admitted as a general
principle, it is perhaps through a study of animal behaviour
that we are best able to realize its full range and extent.
Biologists are so largely, and quite wisely, occupied in the
study of morphological and physiological problems, which
admit of a treatment more exact than the most ardent
advocate of the investigation of behaviour, under natural
or even under experimental conditions, can claim ; they devote,
again quite rightly, so large a share of attention to the
variation and natural selection of adaptive structure in its
adult condition and embryonic stages; the pendulum of
opinion has, under the teaching of Professor Weismann,
swung so far in the direction of the non-acceptance of the
hereditary transmission of characters individually acquired

THE BIOLOGICAL ASPECT 311

through intelligent adjustment or otherwise; that the part
played by consciousness in the evolution of the higher and
more active animals is apt to pass unnoticed or unrecorded.
It is well, therefore, to put in a reminder that a great number
of animals would never reach the adult state in which they
pass into the hands of the comparative anatomist save for the
acquisition of experience, and the effective use of the conscious-
ness to which they are heirs ; that their survival is due, not
only to their possession of certain structures and organs, but,
every whit as much, to the practical use to which these posses-
sions are put in the give and take of active life; and that
many interesting problems which are keenly discussed by
evolutionists in the light of natural selection presuppose
conscious situations which are more or less tacitly taken for
granted.

Let us cast a rapid glance over some of these topics
of biological discussion. The fascinating subject of mimicry,
involving as it necessarily does the discussion of the value
of warning colours and behaviour, a subject opening up an
extensive group of problems so brilliantly studied by Professor
Poulton, is meaningless save in so far as there is implied a
conscious reaction to colour and form on the part of animals
which can learn from experience. The warning colours re-
instate a conscious situation, so that, misled by appearances, a
bird mistakes the mimicking insect for its nauscous “ model.”

The whole range of behaviour, included under play,
experimentation, and practice, on the importance of which,
following the lead so ably given by Professor Groos, we have
insisted, is equally meaningless, save as a means to the acquisi-
tion of serviceable experience for use in the more serious
business of after-life; and experience is the establishment,
through association and coalescence, of conscious situations
which possess guiding value. And if, as we shall hereafter see,
they may also be regarded as a means of securing pleasure,
as a psychological end of behaviour, it is not less obvious that
it is only through the development of consciousness that such
a psychological end can have any existence.

312 THE EVOLUTION OF ANIMAL BEHAVIOUR

It matters not if the particular form assumed by play and
experimentation be largely dependent on instinctive tendencies.
For all the phenomena of instinct, profoundly organic as are
the modes of behaviour comprised under this head, definite as
are the inherited co-ordinations in the most typical examples
of its occurrence, have also, except in some doubtful cases, a
conscious aspect. At any rate this is the case in so far as
instinctive response forms the hereditary basis on which is
reared a more nicely adjusted intelligent edifice, in so far
as instinctive procedure is subsequently modified and guided
by acquired experience, in so far as there creeps in that
“little dose of judgment’ which Huber found in bees, Lord
Avebury attributes to ants, Dr. Peckham sees in spiders and
solitary wasps, and all observers find in birds and mammals.
For if in these cases instinctive behaviour were unconscious,
it would, as such, remain outside experience ; and if outside
experience, there could be no data on which consciousness
could base any modification of inherited behaviour, no
opportunity of taking up the ready-formed responses into the
mental synthesis and utilizing them for the wider ends of
intelligent purpose.

In social behaviour there is a reciprocity of suggestion
between the members of the community. And such sugges-
tion is operative through an appeal to consciousness. How-
ever instinctive the forms of procedure may be in social
insects, there remains much beyond which is hard to explain
on the hypothesis that there is, in them, nothing analogous to
a conscious situation ; while in such vertebrates as birds and
mammals we cannot but believe that consciousness is the main
determinant of much behaviour which scems to imply the
germs, or more than the germs, of sympathy. The little
monkey I saw in Hamburg cuddling up caressingly to a
wounded companion, must surely have experienced a conscious
situation analogous to that which prompts a child to nestle
alongside her companion in distress. And he who has seen no
signs of sympathy in dogs, has either watched their behaviour
in vain, or is himself lacking in sympathy.

THE BIOLOGICAL ASPECT 313

In sexual selection by preferential mating, even if we follow
Professor Groos in believing that it isa special mode of natural
selection, the conscious situation is essential. If we accept the
theory in any form, we must regard the adornments, antics,
and display of the male as an appeal in some way to the
consciousness of the female, whatever particular form the
effects in that consciousness may take, whether the appeal
evoke a sense of beauty, or simply be a means of exciting to
the consummation of the natural end of courtship. Even if
we follow Mr. Wallace in regarding plume and song as “ re-
cognition marks,” it is only by their appeal to consciousness in
this way, if in no other, that they are of any biological value.
And this, of course, applies equally to the whole range of his
theory of recognition marks—their sole utility lies in their
being a stimulus to consciousness through which the end of
recognition is secured. So, too, not only the specialized
behaviour which we dignify by the name of “courtship,” but
every case in which mate is drawn to mate through sight,
smell, hearing—any of the leading senses—testifies to the im-
portance of consciousness in furthering an end of supreme
biological importance.

And if, as Darwin urged, the “law of battle’ among the
males co-operates with preferential mating, as we can hardly
deny, in securing strong, vigorous, and healthy fathers of the
generation they beget, here, too, consciousness is an important
factor. Can we conceive a “law of battle” among unconscious
beings? If success in the combat were a mere matter of brute
strength, it would imply some consciousness in its dull exercise.
But it is more. It is also a trial of skill. Were it not so our
forefathers would not have spent hours in watching a cock-fight,
or laid heavy odds on their particular “fancy.”

We need not labour the theme. In the search for food or
a nesting site, in the capture of prey and escape from enemies,
in all that demands attention, and in all that necessitates
practice, in what M. Houssay calls “the industries of animals,”
and in that which Mr. Hudson calls “ tradition,” consciousness
has a part to play. Even plants unconsciously appeal to the

314 THE EVOLUTION OF ANIMAL BEHAVIOUR

consciousness of insects, birds, and mammals. Their bright,
scented, nectar-bearing flowers, and their sweet, coloured fruits
are means of effecting the biological ends of fertilization and
the dissemination of seeds, but only on condition that their
colours stimulate the sense of sight, and their scent and sweet-
ness the senses of smell and taste. It is, perhaps, going too
far to claim that, wherever sense-organs exist they imply at
least some dim and rudimentary form of conscious situation of
guiding value so far as it goes; for it is possible that in some
cases the coalescence of elementary items of sentience has not
been carried far enough to justify us in speaking of experience
by which the animal can profit. But it is surely not going too
far to claim that, wherever two or three such sense-organs are
gathered together in any living being, there is consciousness in
the midst of them, beginning to exercise that guidance which
serves so markedly to differentiate the typical animal from the
typical plant.

But throughout the animal kingdom, until we reach its
highest development in man, the guidance of consciousness,
important as it is, seems to be almost wholly subservient to a
biological end, that of the preservation of the race, and for the
race of the individual. Practical utility is the touchstone of
animal intelligence, and of the whole range of feeling and
emotion in beings still under examination in the stern school
of natural selection. By this we mean that practical utility has
determined what degree and complexity of intelligence, feeling,
and emotion shall be attained. If the requisite level be not
attained—elimination. Higher levels no doubt bring advan-
tage—-so long as they are practically useful. But in the school
of natural selection useless accomplishments are not much
taught. Although its examinations are in a sense compe-
titive, all are allowed to pass who qualify for survival. But
the competitors become more numerous and the standard for a
pass rises. As the school increases in size higher classes with
harder problems to solve are established. Progress is an inci-
dent of the constant survival of the fittest when there are
variations in fitness,

THE PSYCHOLOGICAL ASPECT 315

II].—Tnre Psycnonogican ASPECT

On the hypothesis of monism, the nature of which, so far
as it bears on our inquiry, was briefly indicated in the fore-
going section, the conscious situation is the psychical or
mental expression of that which for the physiologist is what
we may term a neural situation. As such it does not enter
into the chain of physical causation ; nor do physical events
as such—that is to say, save as experienced—enter into the
chain of mental causation. For mental development they
have no independent existence, and are negligible except in
so far as they enter as items of experience into the conscious
situation.

But altogether apart from the way or ways in which we
may attempt to explain the fact, most of us believe, with
unquestioning confidence, that the growth of practical ex-
perience, somehow associated with nervous changes in the
brain or sensorium, is of real value in the guidance of
behaviour in such manner as to secure biological ends. Con-
scious experience must therefore, in the animal world, serve
its biological purpose, or it will be of no avail. If there be
not a pre-established harmony, there must be an evolved
harmony ; and how such a harmony could be evolved if con-
sciousness be not by some means in vital touch with behaviour,
influenced by and in turn influencing it, we cannot conceive.
The steam-whistle theory of consciousness leaves the matter,
for the evolutionist, in this inconceivable position.

We need not, however, flog a dead horse. We need not
ask how, on the steam-whistle theory, those states of feeling
which we broadly classify as pleasurable could become asso-
ciated with behaviour conducing to welfare, and those which
“we group as hurtful with behaviour which is biologically
harmful. It is more important, again, to notice that, associated
and consonant with the biological end, there arises a psycho-
logical end of behaviour—what we may term, with the quali-
fications before considered,* the getting of pleasure and the

* Vide supra, p. 285.

316 THE EVOLUTION OF ANIMAL BEHAVIOUR

avoidance of pain. This is the purpose of behaviour as
viewed from the psychological aspect. The biological end of
animal conation is racial survival; its psychological end is
individual satisfaction. And the two ends are, in the main
and broadly speaking, consonant—a result which would un-
questionably be secured by natural selection, but is on any
other naturalistic hypothesis difficult of explanation.

But the two ends are not only consonant ; they are supple-
mentary one to the other. During much of the life of the
higher animals there is no need, immediately present and
pressing, for the output of action to meet biological ends.
There are periods of life and intervals of time when the sharp
incidence of the struggle for existence does not call for the
serious business of behaviour. But at these periods and in
these intervals the animal is not inactive ; indeed, it is restless
in its activity. Unless it be weary with unwonted exertion,
or basking in the psychical sunshine of content, due to the
unsought advent of pleasant stimulation or the after-effects
of previous behaviour (for example, when hunger has been
relieved), the healthy animal must be up and doing. This
familiar fact no doubt affords the basis in observation of the
surplus-energy theory of play. But is it necessarily surplus
energy? Is it not rather normal energy which expends itself
in this way when there is no immediate and serious biological
business on hand? And, as Professor Groos has pointed out,
play is seen when we have every reason to suppose there is no
surplus energy, nay, even when the normal energy is at a low
ebb. There is no more pathetic sight than a sick kitten, with
energy obviously much below par, utilizing its little remaining
strength in feeble attempts to play.

It is unnecessary to do more than remind the reader of the
theory elaborated with so much skill and care by Professor
Groos, that the forms assumed by play—in which, it will be
remembered, he includes a very wide range of behaviour—
have a very important indirect biological end in practice and
experimentation. Our present point is, that its direct psycho-
‘logical end is the satisfaction it affords, Without this the

THE PSYCHOLOGICAL ASPECT 317

individual would not be impelled to the continuance of per-
formances which occupy a wide space in the field of animal
behaviour in which the biological end has reference, not to
present requirements, but to future needs.

No one has given better expression to the sway of this
psychological end than Mr. W. H. Hudson. “We see,” he
says,” “that the inferior animals, when the conditions of life
are favourable, are subject to periodical fits of gladness, affect-
ing them powerfully and standing out in vivid contrast to
their ordinary temper. And we know what this feeling is—
this periodic intense elation which even civilized man occa-
sionally experiences when in perfect health, more especially when
young. There are moments when he is mad with joy, when
he cannot keep still, when his impulse is to sing and shout
aloud and laugh at nothing, to run and leap and exert himself
in some extravagant way. Among the heavier mammalians
the feeling is manifested in loud noises, bellowings, and
screamings, and in lumbering, uncouth motions—throwing up
the heels, pretended panics, and ponderous mock battles.

“In smaller and livelier animals, with greater celerity and
certitude in their motions, the feeling shows itself in more
regular and often in more complex ways. Thus Felidee, when
young, and in very agile sprightly species, like the puma,
throughout life, simulate all the actions of an animal hunting
its prey—sudden, intense excitement of discovery, concealment,
gradual advance, masked by intervening objects, with intervals
of watching, when they crouch motionless, the eyes flashing
and tail waved from side to side; finally, the rush and spring,
when the playfellow is captured, rolled over on his back, and
worried to imaginary death. Other species of the most
diverse kinds, in which voice is greatly developed, join in
noisy concerts and choruses; many of the cats may be
mentioned, also dogs and foxes, capybaras and other loquacious
rodents; and in the howling monkeys this kind of perform-
ance rises to the sublime uproar of the tropical forest at
eventide.

* “ Naturalist in La Plata,” pp. 280, 281.

318 THE EVOLUTION OF ANIMAL BEHAVIOUR

“Birds are more subject to this universal joyous instinct
than mammals, and there are times when some species are
constantly overflowing with it ; and as they are so much freer
than mammals, more buoyant and graceful in action, more
loquacious, and have voices so much finer, their gladness
shows itself in a greater variety of ways, with more regular
and beautiful motions, and with melody. But every species
or group of species has its own inherited form or style of
performance ; and however rude and irregular this may be,
as in the case of the pretended stampedes and fights of wild
cattle, that is the form in which the feeling will always be
expressed.”

That all this, which Mr. Hudson so graphically describes,
belongs to the psychological aspect of animal behaviour and
is directly prompted by conative tendéncies whose immediate
end is conscious satisfaction, the mere joy of unrestrained and
healthy activity, may be freely admitted, without denying that
all this exuberant psychical life owes its evolution to the fact
that it is in consonance with and supplemental to biological
ends which secure survival. It is with animals as it is with
man; play is the preparation for earnest. As I have else-
where said,* what our national games have done for the
English race it is difficult to overestimate. They train us to
use our bodies and expend our energies to the best advantage.
An old soldier, watching a football match, said, “'That’s the
training for our future soldiers and sailors.” The playing
fields are the finest school of organized co-operation in the
world. But, apart from compulsion, a boy will not enter into
the game with that zest through which alone it acquires real
value for training, unless there be an immediate psychological
end in the satisfaction he derives. And with animals practice
and preparation for the business of life could not occur if the
ultimate biological purpose of it all were not supplemented by
the enjoyment it brings for its own sake.

But in animal play, as indeed in that of human youth, we
are perhaps a little apt, in laying stress on the bodily skill and

* « Psychology for Teachers,” p. 70.

THE PSYCHOLOGICAL ASPECT 319

readiness of response to which it so effectually ministers, to
forget that it is also a psychological training. In technical
phraseology, we are disposed to fix our attention on the acquired
co-ordination of act and movement rather than on the correla-
tion of conscious data, which renders possible the skilful per-
formance. And yet, rightly considered, the behaviour itself is
simply the outcome of a conscious situation, duly elaborated,
and knit together through the association and coalescence of
its constituent data. It isa means to the unification of con-
sciousness by bringing into relation scattered and, at first, quasi-
independent sensory and emotional elements. Success is only
attained through the concentration of attention and effort on
that which is the centre of interest and also the focus of
endeavour. And this close attention and well-directed effort,
which are trained in the playful output of energy, are just the
mental qualities which will stand the animal in good stead
when the real incidence of life’s struggle comes upon it, when
the reward of success is survival and the penalty of failure
elimination. For they are not merely physical qualities, though
their effects are bodily movements of attack and defence, of
active escape, or merely “lying low.” They are essential
psychological features of a unified and well-directed conative
process.

In the fairly abundant play-time of animal life, this unifica-
tion and direction of conative process can take form under
conditions wherein the preliminary failures which accompany
all forms of learning do not entail the severe penalty of
elimination. If we may so put it, and so apply a deeply
instructive parable, Natural Selection says to her more favoured
children, in which conscious situations can be developed, “‘ Here
are the talents with which I have endowed you; make use of
them till I come, as come I shall in due time.” This animal
puts them out to usury in play ; that animal keeps them laid
up in the napkin of inactivity. Then Natural Selection, the
austere one, comes ; gives the commendation of survival to the
animal that had learnt to put its talents to use in the period of
preparation, and condemns to elimination that which had not

320 THE EVOLUTION OF ANIMAL BEHAVIOUR

traded with his talents at the bank of play. In animal life, on
the perceptual plane, we have the same need for training in little
things and seemingly unimportant matters in preparation for
the stress and storm which may, nay must, come upon them, that
we find in men and women on the higher ethical plane. To
those who think that the play of animals is too trifling a thing
to affect the question of survival, we would suggest the applica-
tion, with a necessary difference, of the thought which Miss
Edith Simcox puts into the following words: ‘“‘ Does it,” she
says, “seem a trifling thing to say that in the hours of
passionate trial and temptation a man can have no better help
than his own past ? Every generous feeling that has not been
crushed, every wholesome impulse that has been followed,
every just perception, every habit of unselfish action, will be
present in the background to guide or to sustain. It is too
late, when the storm has burst, to provide our craft with
rigging fit to weather it ; but we may find a purpose for the
years that oppress us by their dull calm, if we elect to spend
them in laying up stores of strength and wisdom and emotional
prejudices of a goodly human kind, whereby, if need arises, we
may be able to resist hereafter the gusts of passion that might
else bear us out of the straightforward course.” To apply the
thought, the trifles of play supply the psychological rigging
which alone can save the animal craft in the coming storm of
the struggle for existence. And the point on which we have
to lay special stress in this section is, that it is psychological
riggine—or, if this seems to lay too much emphasis on the
genesis of conscious situations, we may at least urge that the
psychological ropes are of co-ordinate importance with the bio-
logical spars.

So far, then, we reach the following conclusion : that if we
classify the behaviour of the higller and more intelligent
animals under two heads, the one comprising all those acts
which are of direct biological value in enabling the animal to
escape elimination under the immediate stress of the struggle
for existence, and the other including all those acts which are
of indirect preparatory or educative value, the latter, which are

THE PSYCHOLOGICAL ASPECT 331

under their biological aspect not less important than the former,
are under their psychological aspect of perhaps even greater
importance. For the conditions of actual struggle are not
those under which mental development could most easily be
furthered, though they are those in which it is most effectually
tested. Hence, the more intelligent animals pass through a
period during which they are more or less shielded from the
incidence of natural selection by their parents, and this is the
period of play and of psychological education. And the tendency
to play is so far organic, in that it is dependent on inherited
instinctive propensities, and so far psychological in that it is
accompanied by a felt want, which constitutes a conative
impulse finding its appropriate end in the consciousness of
satisfaction. But play—if we accept the term as the group-
name for all those modes of behaviour which fall under our
‘second class, those of indirect biological value—does not cease
with the period of youth ; it occupies all the intervals in the
more serious business of animal life. And no discussion of animal
behaviour can be adequate which does not assign to this class its
due place, alike in biological and in psychological evolution.

The whole value of experience lies in the linkage and
coalescence of the data afforded to consciousness. It is true
that an inherited nervous system supplies the organic con-
ditions of that physiological linkage and functional coalescence
of which experience is the psychological expression. It is true
that this physical integration secures a ready-made grouping of
the conscious data which are the concomitants of orderly mole-
cular changes in the brain or analogous sensorium. Still, it also
remains true that the value of experience lies in the further
linkage and coalescence that is acquired by the individual in
the course of what we may fitly call its education. Every step
in this education gets its psychological sanction through the
satisfaction it affords in consciousness; and the time of
acquisition is not during the stress of examination in the
actual struggle for existence, but rather in the youthful period
and in the subsequent intervals of preparation and practice,
during the play-time of animal life.

: ¥

322 THE EVOLUTION OF ANIMAL BEHAVIOUR

The examination analogy—if, indeed, it may not be rightly
regarded as something more than an analogy—may be pressed
a little further as a means of fixing our attention on two points
which are worthy of consideration. The first is that, in the
preparation for the examination, specifie practice as much of
it is, cramming is not the system exemplified by the higher
animals. A good all-round education in the acquisition of
conscious situations more or less coalescent into a unified
system of experience, and in their effective utilization without
unnecessary delay and bungling along more or less converging
lines of practical behaviour ; this is what secures a “ pass”
in survival, especially where the circumstances of life have
reached a considerable degree of complexity. The instinctive
act, with its relatively definite response to a question which is
almost certain to be set to every candidate for survival, is that
which is the analogue in behaviour to the result of a system
of cram. Organic nature does employ this system in the lower
classes of her school ; definite responses are ground into merely
instinctive types generation after generation, and the right
answers are given, automatically and unintelligently, whenever
the oft-recurrent questions are set. But this will not do when’
the questions require the exercise of intelligence, when they
are of the nature of problems, with just those delicate but not
unimportant shades of difference which baffle the candidate
who has been drilled in a merely mechanical fashion. Hence
the cramming of instinct does not suffice for animals whose
environment presents problems of greater variety and greater
complexity. Intelligence is required to meet the particular
combinations as they arise. The greyhound, which is loosed
on a hare, has never seen that hare run in exactly that way
over that special tract of country. But he has been trained
in such situations, and is thus prepared to meet the special
problem in its details as they present themselves in the light
of the experience he has gained of other like problems. And
his skill in pursuit has not only been gained through education
in coursing. In a thousand ways, as puppy and dog, he has
learnt how to use well those sinewy limbs. The training of

THE PSYCHOLOGICAL ASPECT 323

his whole life is brought to bear on the question immediately -
before him.

The general bearing of these facts is obvious. Play, as a
means of animal education, is varied, and has for its end all-
round training of the animal mind in its sphere of operation.
Although there are some specific propensities, certain observ-
able trends of behaviour, as in hunting-play, courtship-play,
and the like, we must not expect, nor do we find, anything
like stereotyped definiteness of conative activity. We find that
freedom and elasticity in animal education which is, perhaps,
more often advocated than carried into practice in human
education.

The second point arising out of the examination analogy
is, that its range determines the level of preparation therefor.
It is, for animals, a practical examination, not a theoretical.
Not a single question is set demanding an explanation. The
problems are such as can be solved by intelligence, not such
as require the exercise of reason, as we have used the term in
foregoing pages. These higher problems are only set when
the sixth form is reached, and there is no conclusive evidence
that any animals get into the sixth. This, however, is entirely
a question of evidence, and many of us will be glad to welcome
them there, if proved ability to deal reflectively with ideational
questions justifies their promotion.

If any of them do belong to this form, they have probably
got there through play. For in the stress of the actual exa-
mination there is not much time for reflection. Or perhaps
‘we may rather say that, not in actual struggle, and not in
‘active preparation for it in play-time, but in intervals of
leisure between both, when the animal lies quietly turning
over in his mind we know not what, will experience be re-
viewed, and generalizations drawn as to the why of events in
this strange world. Probably the animal accepts things as
they are, and does not trouble about their explanation. But
it may not be so. At any rate, if animals lack the means of
descriptive inter-communication, and have no words as con-
crete pegs on which to hang abstract ideas, their explanations

324 THE EVOLUTION OF ANIMAL BEHAVIOUR

cannot be carried far. Theories without the power of dis-
putation would be a poor solace in leisure moments.

One more point may be noticed with regard to the psycho-
logical aspect of the evolution of behaviour—the reciprocal
action of intelligence. It is the intelligence of others that
introduces so much variety and complexity into the environ-
ment. Hunters and hunted, combatants, rivals, mate and
mate, enemies or companions in their varied aspects, introduce
through their intelligence complications which only intelligence
can meet. And, as intelligence begets intelligence, so do
emotional attitudes beget answering emotional states. Psycho-
logical evolution translated into practical behaviour gives rise
to situations of reciprocal complexity. This point of view is,
however, so familiar, that nothing need be said in its further
elucidation. The behaviour of any given animal does not
stand alone, but is closely related with the behaviour of others.
Among social animals the relationships are peculiarly close,
and it is among them that the psychological aspect of behaviour
reaches its highest expression.

IV.—Continurry in Evo.urion

Under the head of organic behaviour, in the widest ac-
ceptation of the term, fall the whole of physiology, the whole
of embryological development, nay, more, the whole of organic
evolution ; while niental evolution, in all its stages, may be
regarded as the psychological aspect of that which, from the
physiological aspect, is the evolution of nervous systems. Life
itself is the behaviour of a particular kind of substance which
is found more or less abundantly under natural conditions.
No other known substance behaves in this way, and so ignorant
are we as to the conditions of its natural origin, that it is
useless to guess at a scientific explanation. And even if we
knew all the antecedents and conditions of its origin we should
be no nearer a comprehension of why protoplasm has the
peculiar properties which we find it to possess. That is a
question to which science can give no answer. Who knows

CONTINUITY IN EVOLUTION 325

why a certain compound of oxygen and hydrogen in certain
proportions has the properties of that which we call water ?

Let us note the distinction between saying, as we said
above, that life is the behaviour of protoplasm, and asserting
that life is the cause of this behaviour. The one is a scientific
statement of observed fact, the other an explanation of the
fact in metaphysical terms, a reference of the fact to its under-
lying cause. So long as we quite clearly understand that we
are talking the language of metaphysics, we may speak of life
as a cause of organic behaviour; but let us be careful to
remember that the statement has no more value for science ;
than the assertion that aqueosity is the cause of the behaviour '
of water.

Leaving on one side, then, the natural origin of protoplasm,
the conditions of which are unknown, we find that, as a matter
of observation, every bit of living substance, the history of
which has been traced, is a fragment detached from some other
bit which behaved in the same way. This is the basal fact of
the continuity of organic evolution. But such a detached
fragment has the property of increasing by taking up from
the environment more of those elementary materials from
which it is itself compounded in subtle synthesis. Nay,
further, every fragment of which we know the history is found
to increase in such a way as to reach, in form, structure, and
idiosyncracies of behaviour, the likeness of the organism—
plant or animal—from which it was derived. In the higher
plants and animals the separated fragments or cells are the
ova and sperms, or their equivalents, which unite, with fusion
or coalescence of their nuclear matter, and thus give rise to a
new individual in the course of embryological development.

Now, as we have already seen, much modern biological dis-
cussion centres round the question whether the detached repro-
ductive fragment, ovum or sperm as the case may be, is derived
from the whole body of the parent, by what Darwin termed
pangenesis or in some other way, or only from germinal sub-
stance set apart in development for this end. And we have
provisionally accepted the hypothesis that it is the direct

326 THE EVOLUTION OF ANIMAL BEHAVIOUR

descendant of other reproductive cells ; and that, throughout
a long ancestry, stretching back into the far past, there never
occurs in the direct line of genealogical sequence, any highly
differentiated, cell, such as a gland-cell, muscle-cell, nerve-cell ;
never, with certain reservations into which we need not enter,
is found the representative of any tissue save that to which the
reproductive function is restricted., In technical phraseology,
the continuity of organic evolution is due to the continuity of
the germinal substance.

During embryological development the fertilized ovum—

‘consisting of two fused fragments of this germinal substance—
gives rise to a host of ordered and marshalled cells, which are
divisible into two groups: the one forms the body with ‘its
muscles, bones, glands, digestive system, skin, sense-organs,
nerve-centres, and so forth ; the other forms,a reserve store of ©
germinal substance, from which are derived the ova and sperms.
The former take no direct share in reproduction ; they are off
the line of continuous descent ; they die without issue. But
they protect and minister to the reproductive function of the
second group—the potential ancestors of the races to follow.
But all instinctive and intelligent behaviour is the outcome of
the orderly working of the nervous system, is initiated through
sensory stimulation, and is executed by the motor organs ; and
all the structural parts, through which such behaviour is pos-
sible, belong to the body—that which dies without issue.
How, then, can instinct and intelligence be inherited? In a
sense they are not inherited. The nervous system which is
their organic basis begets no heirs. But it is begotten of
germinal substance, which not only produced the body of
which the nervous system is a part, but also handed on, with
that body, samples of the same germinal substance capable of
reproducing a similar body and a like nervous system. Herein
lies the basis of heredity.

The stress of the struggle for existence falls upon the
body ; and instinctive or intelligent behaviour is a means to
its preservation in the struggle for existence. According ag
it survives or not, will the samples of germinal substance it

CONTINUITY IN EVOLUTION 327

contains fulfil their biological end or perish with it. Natural
selection secures the survival of those animals which bear the
* seed from which their like will be developed.

On this view all variation arises within the germinal sub-
stance, but it is manifested in the body which is its product.
How variations arise we do not know with any exactness of
detail. That the germinal substance is influenced in its nutri-
tion and in other ways by the surrounding tissues is highly
probable; and this influence may lead to changes which are
the source of variations ; but it is very doubtful whether such
influence can be what we before termed “ homeeopathic.” * It
is improbable that the formation of the nerve-connections
involved in intelligent. behaviour which has grown habitual
through repetition, can so influence the germinalells as to
give rise to variations of like nature. In other words, acquired
habit is probably. not a direct determinant of an inherited
variation of like nature in instinctive behaviour. Apart fram
such influence the only source of variations which can be
assigned is either the differential division of nuclei in prepara-
tion for the process of fertilization,t or the process of fertiliza-
tion itself. The union of perhaps differentiated germinal
substance from two distinct parents affords the opportunities
for the admixture and compounding of hereditary qualities in
the two samples, from which variations favourable or the
reverse may arise.

It is now generally recognized, however, that the origin of
variations is a problem quite distinct from that of the survival
of those whose direction is favourable to that end. The theory
of natural selection, as such, does not pretend to offer any
explanation of the manner in which variations arise ; though
of course a complete theory of organic evolution must assign
the antecedents and conditions of organic progress in all its
varied phases. We know that variations do occur ; we know,
too, that more individuals are born than survive to procreate
their kind ; and, on the theory of natural selection, we draw
from these data the conclusion that, on the average, the

* Supra, p. 36. + Supra, p. 13.

328 THE EVOLUTION OF ANIMAL BEHAVIOUR

animals that escape elimination are those in which the varia-
tions are of such a nature as to conduce to this end.

It will be seen that, on the hypothesis of organic heredity,
thus briefly sketched, continuity can, in strictness and, as we
may phrase it, in its first intent, only be predicated of the
-germinal substance ; but that this substance gives rise to pro-
ducts—active vigorous animals behaving in certain ways, each
after his kind—which hold similar germinal substance in trust
for future use. Natural selection deals with the trustees ; and
if they succumb, that which they hold in trust is lost. To put
the matter in another way : Nature says to the germinal sub-
stance, “By your products you must be judged in accord-
ance with the criterion of utility and efficiency.” Practical
use in the give and take of active life is the touchstone of all
behaviour which makes for survival. This being secured, there
may be a balance of behaviour for other purposes. But in
animals the balance is not of large amount, and other purposes
have not taken form and direction. Jt should be clearly
noticed that, on the hypothesis we are considering, use is
the test of survival, and though it is not the direct cause
of variations, it affords their sanction in survival. That
animal escapes elimination whose behaviour is of practical use ;
and it holds in trust for the future a store of germinal sub-
stance from which is produced a successor capable of behaving
in like manner.

The whole drama of organic evolution may be regarded as
the realization in a succession of individuals of the evolving
potentiality of continuous lines of germinal substance. The
successive individuals die—but the germinal substance lives
on in their heirs, if they have any. In virtue of what inti-
mate and hidden structure or disposition of parts the germ
possesses this potentiality we do not know. The ovum of
a dog is a microscopic speck less than one-hundredth of an
inch in diameter ; the sperm is far more minute. They unite,
and their nuclei coalesce. The cellular product divides and
subdivides. The cell colony absorbs nutriment from the
maternal tissues, Division proceeds apace, and the cells are

CONTINUITY IN EVOLUTION 329

marshalled and ordered in embryological development ; definite
tissues are formed ; the stages of their genesis can be pre-
dicted with accuracy ; and in due time a puppy is born which
shall grow to the likeness of its parents and behave as they
behaved. We can trace the succession of events; we see
that they form a related series; we have good reason for
believing that the state of matters at any one moment is the
antecedent condition of the state of matters at the succeeding
moment. More than this science cannot say. The underlying
cause is, for science, hidden in the mists of the unknown. Even
for metaphysics it is but part of the force that beats through
the universe and makes it not a chaos but a cosmos—a force
known to us only in its effects.

It will thus be seen that the conception of continuity in
organic evolution has, broadly considered, a threefold aspect.
First, there is the continuity of the germinal substance through
whose reproductive behaviour under the appropriate conditions
embryological development occurs; secondly, there is con-
tinuity in this. embryological development, stage by stage,
from the fertilized ovum to the adult which is its final product
and expression ; thirdly, there is continuity in these final pro-
ducts, in the animals whose organic, instinctive, and intelli-
gent behaviour lie open to our study and investigation. The
first is germinal, the second developmental, the third evolu-
tional continuity.

Before attempting to summarize some of the contributions
afforded by our inquiry towards the doctrine of continuity in
the last of these three aspects, we must pause for a moment to
consider how far and in what sense continuity can be predi-
cated of mental development.

We have regarded the conscious situation as the psychical
aspect of a nerve-situation in the sensorium ; and the nervous
system, capable of behaving in this way, is in developmental
continuity with the germinal substance of the fertilized ovum.
But what shall we say with regard to the psychical aspect ?
Two hypotheses seem open to us, each of which presents diffi-
culties, but of different kinds. The first is, that when the

330 THE EVOLUTION OF ANIMAL BEHAVIOUR

organic development of the nervous system reaches a certain
level and order of complexity consciousness emerges, how and
whence we know not. The second is, that consciousness is
developed from sentience, which is the concomitant of all
organic behaviour ; which accompanies life wherever it occurs,
and therefore shares the continuity of the germinal substance.

The difficulty inseparable from the first hypothesis, is that
it is contrary to the analogy of all that we know or infer else-
where throughout the realm of nature. Huxley * likened its
emergence to the production of heat when an iron bar is
struck by repeated blows of the hammer. But this analogy
will not hold ; for heat is a mode of energy, and only emerges
through the transformation of other and pre-existing modes
of energy. A certain amount of the energy of motion in the
massive hammer-head is transferred to the iron rod, and
assumes the form of that molecular vibration which we call
heat. And by what amount the one is the gainer, by that
amount is the other the loser. But we have no reason to
suppose that the like takes place in the origin of the mental
concomitants of neural changes. No portion of the brain’s
store of physical energy is drained off to form the rivulet of
consciousness. Now, whenever we speak of a product else-
where in nature, we mean a specialized bit of something pre-
existent. Water is the product of pre-existing oxygen and
hydrogen. Heat is the product of other forms of energy.
But this is not so on the first hypothesis, according to which
consciousness emerges when the functional activity of the
nervous system reaches a certain level and order of complexity.
The mental concomitants are not “ products,” in the recog-
nized sense of the term. Furthermore, although on this
hypothesis we may still speak of what was termed above
evolutional continuity in the mental concomitants, there is
nothing analogous to either developmental or germinal con-
tinuity.

On the second hypothesis, according to which sentience is
the concomitant of all organic behaviour, such developmental

* « Collected Essays,” vol. i., p, 239.

CONTINUITY IN EVOLUTION. 331

and germinal continuity, or their analogues in the psychical
order of being, are rendered conceivable. Consciousness is
regarded as a developed form of sentience. But the sentience
is wholly hypothetical. It is at best a “may be,” and its
existence is incapable of proof. And science is rightly im-
patient of hypotheses the validity of which .cannot in any way
be verified. Our safest course, therefore, is to accept that
which is common to both hypotheses, evolutional continuity,
and for the rest to be content with a confession of ignorance.

We have already drawn attention to the fact that mere
sentience, if it exists, has no power of guidance over organic
behaviour ; but consciousness, when it emerges, is a concomi-
tant of nervous processes which determine the nature and
direction of such nerve-changes as are the antecedents of
intelligent behaviour. The steps by which this control is
established are unknown. It is, indeed, probable that con-
scious guidance arises as an accompaniment of the differentia-
tion of controlling centres from the automatic centres of the
nervous system ; but of how this takes place we are as igno-
rant as we are of many other differentiations in the course
of embryological development and evolutional progress. Of
those nervous arrangements within the brain which are the
physiological concomitants of the far later mental processes of
reflection, abstraction, generalization, and the formation of
ideals, we are, if it be possible, even yet more profoundly
ignorant. Nor would it serve any good purpose to indulge in
speculation where there are not even the data to enable us so
much as to hazard a probable guess. The utmost we are
justified in attempting is to show how organic behaviour leads
up to and affords the requisite data for the exercise of intelli-
gence, and how both supply the necessary preliminary stages
in the development and evolution of what, following Dr.
Stout, we have termed ideational process. This we have
endeavoured to do in preceding pages; and all that is now
required is to conclude our inquiry with a brief summary by
which the results, as affording some basis for evolutional con-
tinuity, may be focussed. 2

332, THE EVOLUTION OF ANIMAL BEHAVIOUR

We regard reflex action and instinctive behaviour, broadly
considered, as genetically prior to that which is intelligent.
Their development in the individual and their evolution in
the race are reached by the differentiation and integration of
nerve-centres. In the abdominal region of the crayfish, for
example, special centres are differentiated for the behaviour
of each pair of swimmerets; but these are so integrated that
the whole series of like abdominal appendages swing rhythmi-
cally with co-ordinated movements. Now, when a sensorium
is developed, it does not have to group by an act of con-
scious selection and deliberate arrangement the multiplicity of
scattered sensory data which it receives ; it does not have to
organize from diverse and hitherto unrelated elements some
sort of system in experience: it receives them as a physio-
logical heritage already grouped, and to some extent organized.
Stimulus and response are organically linked ; and within the
response inherited co-ordinations, often exceedingly complex,
afford a correlated group of sensory data. Just in so far as
organic heredity has provided a working system of bodily
parts, does consciousness receive systematic information of
their orderly working. No doubt it is true that the develop-
ment and evolution of the sensorium proceeds pari passu with
the development and evolution of reflex actions compounded
and co-ordinated to give rise to instinctive behaviour. No
doubt the progress of the one is in close touch and relation
with the progress of the other; for such relation receives the
emphatic sanction of utility. Still it is none the less true
that in individual development, as in racial evolution, the
organic takes the lead. What is intelligently acquired is
something added to that which has been engrained, through
natural selection or otherwise, as a potentiality of the germinal
substance. What we have first to note, then, is that organic
evolution provides ready-grouped data to consciousness.

The second point is, that the germs of abstraction and
generalization, or rather processes which are the precursors of
abstraction and generalization, arise, and cannot fail to arise,
in the genesis of experience from the performance of inherited

CONTINUITY IN EVOLUTION 333

responses, and from the coalescence of their results into a
conscious situation. To a quite young chick I gave pieces of
yellow orange peel, which were found to be distasteful and
rejected. In Dr. Stout’s phraseology, they acquired meaning
in experience. Can one doubt that the colour and taste were
thus rendered predominant, and that the shape, size, and other
qualities of the bits of orange peel remained practically un-
noticed ? Shortly afterwards the chick was given chopped
and crumbled egg ; the fragments of “ white” were eaten, but
the bits of hard-boiled yolk were untouched. They possessed
a sufficient general resemblance to the orange peel to carry the
same meaning. In many ways particular qualities of objects
are emphasized in so far as they incite to behaviour ; they form
centres of biological interest, just as the abstract quality of
ideational thought is the centre of rational interest on a higher
plane of mental development. And in many ways objects present-
ing certain salient features in common, amid differences which
remain unnoticed, are unconsciously grouped as the starting-
points of similar perceptual situations, just as in the generaliza-
tion of ideational thought similar relationships are deliberately
grouped as the starting-points of like conceptual situations.
Both are purposive and have an end, which we as investigators
are able to assign; but only for reflection and conceptual
thought are they also purposeful—the end being foreseen and
realized, not only by the investigators, but by the agent con-
cerned. And the purpose or end itself is in the two cases
different. In the one case it is the biological end of practical
behaviour ; in the other case it is the rational end of explana-
tion—abstraction and generalization being deliberately used as
a means to this latter end. The question has again and again
been asked : Do animals reason? And different answers are
given by those who are substantially in agreement as to the
facts and their interpretation, but are not in agreement as to
their use of the word “reason.” Perhaps, if the question
assume the form—Are animals capable of explaining their own
acts and the causes of phenomena ?—the position of those who
find the evidence of their doing so insufficient may be placed in

334 THE EVOLUTION OF ANIMAL BEHAVIOUR

a clearer light. This is what is generally meant by the state-
ment that animals have probably not reached the level of
rational beings.

But even if they have not reached this level, their perceptual
processes supply the antecedent conditions which are necessary
if this level is to be attained in the course of further evolution.
We have seen that, even in relatively simple cases, where con-
scious situations mark only the beginnings of intelligence, there
is a biological emphasis of some, rather than others, among
what we call the qualities of objects, and there is a grouping,
on biological grounds, of certain things which have some
quality in common—such, for example, as being fit for food.
Here we have at the outset of perceptual development the
germs of processes which are the precursors of the abstraction
and generalization of ideational thought. And in the more
complex conscious situations of the higher animals these pro-
cesses attain to such degree of development as is necessary to
secure more difficult and more remote biological ends, until all
that is necessary, for their rational use, is the quickening touch
of a new purpose, that of explanation.

We have seen that, through what Dr. Stout terms
“manipulation,” and Professor Groos “ experimentation ”—
names applied to a type of behaviour widely exemplified
among the higher animals,—things, as the nuclei of conscious
situations, become differentiated from the environment. One
can hardly question that a fly to the trout, a ball to the kitten,
a bone to the puppy are things distinguished from their
surroundings, and that they become marked off as special
centres of interest. Here on the perceptual plane is a process
which is the antecedent of the conception of quasi-independent
objects on the ideational plane.’ For rational thought the
thing, as object, is not only the centre of a practical
situation leading to behaviour of direct or indirect biological
value, but is the nucleus around which we build all the qualities
which are ascertained by more elaborate manipulation and
experimentation carried out deliberately and of set purpose
for rational ends. It becomes capable of definition with

CONTINUITY IN EVOLUTION 335

the aim of explaining what are its characteristics as an
object.

There can be little doubt that the higher animals become
intimately and practically acquainted with their environment.
The dog who accompanies his master in many a ramble, the
horse who carries him again and again over all the surrounding
country, has a good perceptual knowledge of a somewhat
extended environment. And this, again, is the precursor of the
far more extended conceptual knowledge which leads up at
last to a rational conception of the universe of objects in their
varied relationships. But only through the concentration of
thought rendered possible by much true abstraction. and
generalization,—only through disentangling the relationships
and regrouping them for the purpose of framing an ideal
scheme,—only, in short, by explanation and for the sake of
explanation is this difficult process brought to a more or less
successful issue. ;

Again, there can be little doubt that the higher animals, in
the course of experience begotten of behaviour, reach a per-
ceptual sensing of the bodily self, through experience derived
from the non-projecting senses, in pain and sickness, and
often, we may hope, in the sense of well-being, and the joy of
existence. They do not probably set this self in antithesis to
the not-self. That comes with reflection, and is the result of
ideal construction based on the analysis of experience, with a
view to reaching some explanation of the genesis of experience.
But in their perceptual awareness of the embodied self, they
have that kind of consciousness which affords the necessary
data, for the later conception of the self—when experience is
polarized into its subjective and objective aspects and thus is
explained, so far as science can explain it ; suggesting, indeed,
long ere science has attained this end, metaphysical explana-
tions by reference to underlying causes—too often accepted as
an easy substitute for the difficult tracing out of the antecedent
conditions which science endeavours painfully and by slow
steps to formulate.

It is unnecessary to do more than remind the reader that

336 THE EVOLUTION OF ANIMAL BEHAVIOUR

we have found that such processes as attention and imitation
pass through instinctive and intelligent stages which are the
precursors of the ideational stage, where they reach a higher
expression as deliberately conscious acts. In the young bird
that instinctively pecks at some small, perhaps moving, thing,
which forms the starting point of a piece of responsive behaviour,
we have attention in the germ. When experience has caused the
thing to acquire meaning, attention passes into a succeeding in-
telligent phase ; but only when we desire to explain this meaning,
and attention thus has a deliberate purpose, do we find it entering
upon its higher ideational career. So, too, as we have seen,
imitation is at first a specialized form of instinctive behaviour,
where the response is seen to resemble that which stimulates
it. Later it becomes intelligent when the repetition of the
imitative behaviour is due to the satisfaction it introduces
into the conscious situation. Then, at last, it reaches the
ideational stage, where reflection gives rise to an ideal, which
is to be realized in conduct. The imitation by the child of
its older companions is at first probably intelligent; but
when the child begins to consider why it imitates these and
not those among its companions, he is passing to the ideal
stage, and imitation becomes the sincerest form of hero-
worship. The boy who merely imitates his elder brothers
playing at soldiers because he gets satisfaction from so doing,
becomes the subaltern who has his ideal soldier, and will face
death firmly rather than fall below his conception of how such
a soldier should behave.

We need not again attempt to indicate how among animals
we have the perceptual precursors of the esthetic and ethical
concepts. But we may remind the reader that we endeavoured
to show that intercommunication had its foundation in instinc-
tive sounds; and that it passed into the intelligent stage in
the perceptual life, when these sounds acquired meaning,
and hence became guides to behaviour. This is especially
instructive from our present standpoint, since it is probable
that the passage of communication from the indicating to the
descriptive stage afforded the conditions under which rational

CONTINUITY IN EVOLUTION 337

thought was evolved. For such thought it is essential that
attention should be focussed on the relationships of things.
And no description is possible without making distinctly
present to consciousness these relationships, in time and space,
the data for which are abundantly present in the perceptual
life, though lurking in the background, and needing something
to fix them and to aid consciousness in distinguishing them
clearly. In descriptive communication parts of speech, or
their initial equivalents, afford fixation points for these relation-
ships, and serve to render them distinct. If the reader will
try to describe even the simplest occurrence without introducing
the symbols for the relations which the events bear to each
other, his failure will serve to bring home how essential a
feature this is. In social communication, tlen, we probably
have the key to the -passage from perceptual to ideational
process ; and in this passage description is the antecedent of,
and affords the conditions to, explanation. Words, moreover,
as we have already said, form the pegs upon which we can
hang up, for ready reference, the products of abstraction and
‘generalization, or, to modify the analogy, they form the bodies
of which these products are the rational soul.

If we are ever to trace the passage from the instinctive
through the indicating stage of communication, and so onwards
through the beginnings of description to its higher levels, and
thus to the use of language as a medium of explanation, it
must be through child-study. In every normal human child
the passage does actually take place, though, no doubt, in a
condensed and abbreviated form as an epitomized recapitula-
tion in individual development, of the steps of evolutional
progress. Thus we may obtain a key to the solution of one
of the most difficult problems in evolution by continuous
process—that of the transition from animal behaviour to
human conduct.

INDEX

A

Abstract and general ideas, 57
Abstraction, 166; germs of, 332 ff.
Acceleration, 250

Accommodation defined, 36

Acquired characters, inheritance
of, 35, 110

Acquired instincts (Wundt), 66,
106.

Acquisition defined, 36; ultimately
dependent on natural selection,
289

Adaptation defined, 37

ADDISON on instinct, 63

Aisthetics, animal, 270

Afferent and efferent impulses, 32,
101

Aid, mutual, among animals, 227

Ammophila mode of stinging prey,
75; of carrying prey, 76 ; deposi-
tion of egg, 77; intelligent be-
haviour of, 127

Ameba, 296

Antlers of deer, 15

Ants, behaviour of, 123; inter-
communication of, 198; social
communities of, 205

Aporus, intelligent behaviour of,
126

Appreciation, germs of, 273

Ardour of male in courtship, 269

Argyromeba, instincts of, 79

Arrest of development in egg, 14

Association in coalescent situation,
46

Attention, 242

Avupuson on American night-hawks,
261

338

Avesury, Lord, on ants, 198; on
Van, 200; on aphides and ants,
214; on slave ants, 215; on
intelligence of ants, 218

B

Batpwin, Prof. Mark, on organic
selection, 37 (note), 115; on
functional selection, 163; on
imitation, 179 ff.; ou projective
stage of development, 275

Batesian mimicry, 164

BrEckstTEIN on canaries, 262

Bees, homing of, 131; social com-
munities of, 205

Beetle soliciting food from ant, 213

Bembex mode of carrying prey, 76

Brrue, Dr.,on instinctive behaviour
of ants, 217

Binet, M., on infusoria, 6

Biological value of play, 250; pur-
pose, 294; aspect of animal be-
haviour, 305

Birch-weevil, leafcase of, 121

Birds, instinct of, 84

Bison, behaviour of the, 226

Buackpurn, Mrs. Hugh, on instinct
of cuckoo, 90

ee Fae Dr., on componotus,

Borron on goldfinches’ nests, 136

Bower-bird, observations on, 261,
273

Bucemay, Mr. 8. S., on speech of
children, 203

Buperrr, Mr. John §., on nest-
building, 135

Bullfinch, nest of, 135

INDEX

Cc

Cameron, Myr., on mimetic insects
in ants’ nest, 212

Canaries’ nest, building of, 135

Canon of interpretation, 270

Capacity, innate, 176

Capuchin monkey, imitation in,
188, 278

CakPENTER, W.B., on water-beetle,
299

Catasetum, fertilization of, 29

Cats, Prof. Thorndike’s experiments
on, 147, 184

Causation, idea of, 257

Cell-division in egg, 14

Cerceris, instincts of, 74; locality
studies of, 129

Chalicodoma, parasities of, 78 ;
Fabre’s observations on, 130

Chick swimming, 85; instincts of,
85 ff.; imitation in, 183

Child-study, desirability of, 155,
337

Choice, apparent in Paramecia, 9 ;
in the pairing situation, 266

Ciliary action in Paramecium, 4-10

Circular process (Baldwin), 181

Clepsine, behaviour of, 159

Coalescence in conscious situation,
46

Coincident variations defined, 37;
survival of, 115, 174

Communities, social, of bees and
ants, 205

Companion as centre of special
interest, 244

Componotus, communities of, 210

Conation and impulse, 187, 235

Concept, nature of, 166

Condensation of experience, 162

Conduct implies motive, 60; and
ideal, 278

Congenital responses, 41

Conjugation in Paramecium, 4

Connate instincts, 66, 69

Conscience, ambiguity of
281

Conscious accompaniments of cer-
tain organic changes, 42; aspect
of instructive behaviour, 99

Consciousness, as accompaniment
and as guide, 34; effective, de-

word,

339

fined, 43; as heir to organic
estate, 52; as epiphenomenon,
306

Consentience, 53, 62

Consonance of biological
psychological end, 286, 316

Constancy of environment leads to
stereotyped behaviour, 172

Continuity in evolution, 324; threc-
fold aspect of, 329

Control, the sign of effective con-
sciousness, 43

Co-ordinated acts,69, 100; inherited,
94, 95

Corporate behaviour, 14

Courtship in animals, 259

Coyness of female birds, 2U- ff.

Cranial sense-organs, 301

Crayfish, reflex action in, 298

Creation, special, 297

Criteria of effective consciousness,
43; of intelligence, 120

Cruelty in cat, 277

Cuckoo, instinct of nestling, 90

and

D

Darwin, Charles, fig. of sun-dew
leaf, 26; of Venus’s fly-trap, 27;
of catasetum, 31; on earthworms,
157; on social life of animals, 225;
on human ancestry, 229; on
play and practise, 259; on sexual
selection, 262 ff.; on law of battle,
313

Davis, Prof. Ainsworth, on limpets,
155

Dray, Dr. Bashford, on chick swim-
ming, 85

Deceit in animals, 280

Deferred instincts, 70

Definiteness of instinctive be-
haviour, 66

Description,
terms, 202

involyes relational

_Didunculus, changed habits in,

221
Differentiation and integration of
nerve centres, 167
Disintegration of instincts, 175
Diving, instinctive, 86
Dog, observations on, intelligence
of, 141 ff, 152, 200, 271, 322

340

Duckling, inherited co-ordivation
in, 96
Dytiscus, instinct of, 104

E

Earthworms, Darwin’s observations
on, 157 ff.

Edueation in play, 255, 320 ff.

Effective consciousness detined, 43

Effereut and afferent impulses, 32,
101

Egg, cell-division in, 14

Eimer, Th., on instincts of solitary
wasps, 73; on origin of instincts,
107

Emotions, and feelings, 235 ff. ;
psychological nature of, 246;
evolution of, 282

Energy stored in cell, 23

Equilibrium, tendency to, 296

Eristalis, mimicry of, 164

Esrinas, Prof, on social life of
animals, 230

Ethics, animal, 270

Evolution of organic behaviour, 35;
of consciousness, 61; of instinc-
tive behaviour, 106; of intelligent
behaviour, 155; of social be-
haviour, 225; of feeling and
emotion, 282; of animal be-
haviour, 295; as continuous, 324
ff.

Experience, of value for future
guidance, 44; is it inherited?
48,97; condensation of, 162

Experimentation, 251, 253

Explanation, characteristic of later
phases of mental development,
58, 257

Explosive nature of ccll, 21

Tixpression of emotions, 247

External stimuli to instinctive be-
haviour, 102

F

Fapre on behaviour of Sphex, 77,
171; of Chalicodoma, 78, 129;
of Leucopsis, 79; of Pompilus,
129

Faculty, instinctive, 64

Falcons, training of, 137

INDEX

Fear in birds not inherited in
specific direction, 49, 110

Feelings and emotion, 235 ff.;
evolution of, 282; feeling-tone,
240

Ferns, fertilization of, 24

Fertilization of ferns, 24; of Valis-
neria, 28; of orchids, 29

Fiyy, Mr. Frank, on the acquisition
of experience by young birds, 50

Fission, reproduction of Paramecium

Flight, instinctive, 86

Foren on Componotus, 211

Foster, Sir Michael, on conscious-
ness accompanying reflex action
in pithed frog, 33

Frog, reflex action in, 33, 299, 300

Functional selection, 163

Fungus garden of ants, 216

G

Garner, Mr. R. L., “The Speech
of Monkeys,” 198

Gas-engine, analogy of, 20

General and abstract ideas, 57;
generalization, 166; germs of,
332 ff.

Generic image, 162; situations, 163

Germinal substance, continuity of,
328

GovuLp, Dr., on humming-birds, 273

Green, Mr. E. G., on ants, 210

Greentinch, nest of, 135

Groos, Prof., on instinct, 64; origin
of, 116; on imitation, 187; on
animal play, 248 ff.; on “Love
Play,” 259; on coyness of female
birds, 264; on choice in mating,
267; on make-believe, 280

H

Habits and habitual acts, 107, 177

Have on ants, 199

Hamerton, P. G., on trained dog,
152

Hancock, Dr. Juhn, on cuckoo, 92

Heredity and circumstance, 39;
twofold aspect of, 40; relation of
to use, 170, 176; in evolution as
continuous, 326

INDEX

Homing of bees, 131

Homeeopathic influeice defined, 36

Honey-pot ant, 215

House-martin, nest-building of, 112

Hupson, Mr. W. H., on fear in
birds, 49, 50, 110; on animal
gladness, 317

Hunting play, 254

Hoxtey, T. H., on reflex action in
frog, 300; on consciousnezs as
epiphenomenon, 306 ff.; on con-
sciousness as product of nervous
changes, 330

Hype, Mr., on king crab, 298

Hydractinia, colonial polype, 206

I \

Ideals, distinguish ethics, 278

Ideational stage of mental develop-
ment, 59

Imitation, 179 ff.; three stages of
in child, 192

Impulse io intellizent behaviour,
60; Prof. Thorndike’s use of the
term, 186;
conative process, 235

Independence of automatic and
controlling centres, 43

Infant, congenital responses in, 54

Influence of intelligence on in-
stinct, 168

Inheritance of acquisitiveness, 40

Innate capacity, 176; likes and
dislikes, 119

Insects, instinctive behaviour in,
71; intelligent behaviour in, 123

Instinct, broader and narrower view
of, 99; primary and secondary,
107,109; influence of, intelligence
on, 168; priority of, to intelli-
gence, 173; disintegration of, 175

Instinctive behaviour defined, 63 ;
in insects, 71; in birds, 64; con-
scious aspect of, 98

Integration and differentiation of
nerve centres, 167

Intelligence lapsed, 107; influence
of, on instinct, 168; of ants, Lord
Avebury on, 218; biological im-
portance of, 310

Intelligent behaviour, 117; evolu-
tion of, 155

connection of with |

341

Intelligent process distinguished
from rational, 59, 138

Intercommunication, 193, 336

Interest, 243

Internal factors in instinctive be-
haviour, 102

Irritability, fundamental, property
of protoplasm, 240, 296

J

James, Prof. Wm., theory of emo-
tions, 246, 292

Jays, bathing of, 89; mode of taking
food, 94

JENNER on cuckoo, 92

Junxnines, Dr. H. 8., on behaviour
of Paramecia, 5

K

Kuerner, Dr., on sun-dew, 26; on
sensitive Oxalis, 27; on Valis-
neria, 29

King-crab, reflex action in, 298

Kyicut, Andrew, on Norwegian
ponies, 109

Krovorgine, Prince, on mutual
aid among animals, 227

L

Lamarckian hypothesis, 169, 170,
177

Language, nature of, 195

Lanxuster, Prof, E. Ray, on small-
brained mammal, 168

Lapwing, instinctive behaviour of,
113

Law of battle, 313

Leech, observation on, 159

Leucopsis, instincts of, 79

Lewes, G. H., on lapsed intelli-
gence, 107

Limpets, observations on, 155

Linptey, Dr., on children, 141

Locality, studies by wasps, 128

Locke, John, limitations of animals,
166

M

Macxenzig, Prof. J. §., on ethics,
278; on conscience, 281; on
ambiguity of word“ pleasure,” 285

342

Make-believe, 230

Mammals, early small-brained, 168

Manipulation (Stout), 251

Marcuat, Prof., on instincts of
Cerceris, 74

Mansaatt, Mr. H. R., on instinct,
66

Martin, nest-building of, 112

Marrineav, James, on pleasure
and pain, 284

Mavpas, M., observations on, in-
fusoria, 4

Mayer, Dr. A. G., on mating,
instinct of moths, 83

McCook, Dr., on ants, 214

Meaning, Dr. Stout’s use of term,
46,°243, 268

Mevtioort, Mr. H. B., on behaviour
of wild pigs, 196

Megapodes, instinctive flight of,
87

Meloé, instincts of, 81

Mental development, stages of, 48,
56

Mercier, Dr. Charles, on criteria of
intelligence, 120

Metaphysical explanations, 19;
aspect of instinct, 64; of impulse,
237; of purpose, 294; of will, 307;
of life, 325

Mints, Prof. Wesley, on social
influence on puppy, 220

Miltogramma, parasitic fly, 134

Mimicry, Batesian and Miillerian,
164; intelligent aspect of, 311

Modifiability, 171

Modification, defined, 36; relation
of, to hereditary characters,
169

Mo tter, Herr, on fungus garden of
ants, 216

Monistic hypothesis, 309, 315

Monkey, capuchin, imitation in,
188

Monodontomerus, instincts of, 79

Moor-hen, diving of, 89

Moths mating, instinct of, 83

Motive in rational conduct, 60

Movement plays, 251

Miuuer, Prof. Max, on barrier
between brute and man, 204

Miillerian mimicry, 164, 165

Alystery of life, 18

INDEX

N

Natural selection, shielding of
chicks from, 111; under uniform
and variable circumstances, 175;
in playtime of life, 319

Nervous are, 33; system of higher
animals, 297

Nest-building, observations on, 135

Norns on concept, 166

Norwegian ponies, 109

Nucleus division, 1z

Oo

Object and subject, 245, 276
Octopus, intelligence of, 157
Gicophylla, behaviour of, 210
Orchids, fertilization of, 29
Organic basis of differentiation of
consciousness, 53
Organic behaviour in development,
15
Organic selection, 37 (note), 115
Overproduction of movements, 163
Oxalis, sensitive, behaviour of, 27
Oxybelus, mode of carrying prey, 76

P

Patey, definition of instinct, 64
Paramecium, behaviour of, 3, 296
Partridge, note of young, 93
Prcxuam, Dr. G. W., on instinct,
65; on solitary wasps, 72 ff.,
126 ff.
Pecking instinct of chicks, 93
Peewit, note of young, 93
Pelopzus, instincts of, 72
Perceptual stage of mental develop-
ment, 59
Personality, 245, 257
Pheasants, note of young, 92; in-
herited co-ordination in, 95;
plumage of Argus, 262
Philanthus, prey of, 73; mode of
stinging prey, 74
Physiological aspect
behaviour, 295
Pigeons, nests of, 136
Pigs, wild, behaviour of, 196
Plants, behaviour of, 24

of animal

INDEX

Plastic period of life, 167

Plasticity of tissues, 40; of be-
haviour, 172

Play of animals, 248 ff.; biological
value of, 250; psychological as-
pect of, 256, 311, 316

Prayne, Mr. H. C., on pigeons’
nests, 136

Pleasure, 241; ambiguity in word,
285

Polistes, locality studies of, 131

Pompilus, mode of carrying prey,

76; Fabre’s observation on,
129
Presentative elements distin-

guished from re-presevtative, 46
Primary instincts (Romanes), 107
Projective stage of mental de-

velopment, 275; senses, 304
Pronuba, instinct of, 82
Propensity, instincts as, 64; con-

genital, 177
Protoplasm, fundamental proper-

ties of, 296
Psychological aspect of play, 256;

purpose, 294; aspect of animal

behaviour, 315

R

Rational process distinguished
from intelligent, 59, 138

Reflex action, 31, 35, 298 ff.; rela-
tion of instinct to, 70

Relationships, importance of, 202

Re-presentative elements distin-
guished from presentative, 46

Rhynchites, instinct of, 121

Romangs, G. J., on “ discrimina-
tion” and “perception” in
plants, 32; on instincts of soli-
tary wasps, 73; definition of
instinct, 99; on primary and
secondary instincts, 107, 109; on
ants, 126; on general ideas, 166;
on animal communication, 201;
on cruelty in cat, 277

Romanes, Miss, observations on
capuchin monkey, 188, 278

Roots of spinal nerves, 299

Rorsnsy, Mr. G. A. G., on Indian

ants, 212

343
s

SCHNEIDER on octopus, 157

Scott, Dr. D. H., on fern fertiliza-
tion, 25

Scratching in duckling, 96

Sea anemone, diffused nervous
system of, 32

Secondary ‘instincts
107, 109

Segmental nature of central ner-
vous system, 299

Selection, functional, 163; natural,
shielding of chicks from, 111;
under uniform and variable cir-
cumstances, 175; in playtime of
life, 319; sexual, 261 ff., 313

Self, as ideal construction, 239

Sentience, 62; origin of, 330

Sexual selection, 261 ff., 313

Suarp, Dr. D., on birch-weevil,
121; on Gicophylla, 210

SHEeLiarD, Mr. E. J., observations
on staghound, 144

SHeErRinetTon, Prof., on emotion,
292; on spinal animal, 298 ff.

Shock effects of physiological, 202

Srmcox, Miss Edith, quoted, 320

Sitaris, instincts of, 82

Slave ants, 215

Snails, observations on, 157

Social behaviour, 179; evolution
of, 225

Solitary wasps, instincts of, 72 ff. ;
intelligence of, 126 ff.

Solomon Islands, rats of, 222

Sounds emitted by young birds,
92

Spatpinc, Douglas, on newly
hatched turkeys, 49; on in-
stinct, 99

Special creation, 297

Speech, connection of, with ra-
tional process, 58, 233; so-called,
of monkeys, 198; of children,
203; aids passage from percep-
tual to ideational process, 337

SPENCE on instinct, 63

Spencer, Mr. Herbert, on instinct
and reflex action, 70; on play
due to surplus vigour, 248; on
pleasure and pain, 284, 287; on
survival, 288

(Romanes),

344

Sphex, mode of carrying prey to
nest, 77, 171
Spiders placed in crotch by wasps,

Stereotyped behaviour, 172
Sticklebacks, observations on, 130
Strout, Dr. G. F., on “meaning”
for consciousness, 46, 243, 268 ;
on ideational and perceptual
stages of mental development,
59, 271; on octopus, 157; on
conative process, 235; on the
self, .239; on manipulation, 251 ;
on emotion, 293
Strance, Mr., on bower-bird, 261
Subject and object, 245, 276
Sun-dew leaf, behaviour of, 25
Swimming, instinctive, 85

T

Tarr, Lawson, on begging-cat, 109

Tarpr, M., on imitation as a
social factor, 179

Tendencies, congenital, 177

Tuomas, Mr. Oldfield, on rats of
Solomon Islands, 222

TuornvDIKE, Dr., on swimming of
chick, 85; experiments on in-
telligence, 147 ff.; experiments
on imitation, 179, 183 tf.

Tradition, animal, 220

Trial and error, method of, 139

U

Unity of perceptual process, 240 ;
of biological purpose, 297

Use, super-normal, 169; relation
of, tu heredity, 170, 176

THE

INDEX

vV

Valisneria, fertilization of, 28

Variation defined, 36; origin of,
3827

Venus’s fly-trap, behaviour of, 26

Vigour, play due to surplus, 248

Volition as conative, 238

WwW

Watiace, Dr. A. R., on sexual
selection, 264

WaALuascHEr, Mr., on play as sur-
plus vigour, 248

Wapiti, antlers of, 16

Wasmany, Dr., on insects asso-
ciated with ants, 213

Wasps, solitary, instincts of, 72 ff. ;
intelligence of, 126 ff.

Weir, Mr. Jenner, on canaries,
135

WEISMANN, Prof., on origin of in-
stinct, 108

Wurman, Prof., on Clepsine, 159

Wuitmer, Rey. 8. J., on tooth-
billed pigeon, 221

Will, metaphysics of, 307

Wiuuiston, Dr. 8. W., observation
on Ammophila, 127

Woop, Mr. Foster, on hen-swim-
ming, 86

Worcester, Dr., on megapode, 87

a Prof., on instinct, 65, 99,

Y

Youth, plasticity of, 167
Yucca moth, instincts of, 82

END

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By JOSEPH R. FISHER, B.A,

BarRISTER-AT-Law.
SEeconp EDITION.

Demy 8v0., cloth, 125. 6d.

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Io

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LECTURES ON THEORETICAL AND
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11

PUBLICATIONS OF THE ESSEX HOUSE
PRESS.

These books are printed at Essex House, on the presses used by the
late Mr. William Morris at the Kelmscott Press, which were purchased
by the Guild of Handicraft. Members of Mr. Morris’s staff are also
retained at the Essex House Press, and it is the hope of the Guild of
Handicraft by this means to continue in some measure the tradition of

good printing and fine workmanship which William Morris revived.

A new type is being designed by Mr. C. R. Ashbee, and is nearly
ready ; in the meantime the books of the Essex House Press are being
printed in a fine eighteenth-century Caslon fount, with specially designed
ornaments and initials. It is gratifying to note that the demand for the
limited editions of the books already issued is steady and persistent, and

it will probably not be long before they are out of print.

Subscribers to the complete series of Essex House Publications are
given priority for any new book issued, and the number of subscribers

is constantly increasing.

The following volume of the Vellum Series is just ready :

Keats’ Eve of St. Agnes. The second of the Vellum Series. It
will be rubricated, and will contain a hand-coloured wood-block frontispiece by
REGINALD SAVAGE. Edition limited to 125 copies. Price £2 net.

12

In preparation:

The Psalms of David, according to the text of the Anglican Prayer-
Book.

Erasmus’ Praise of Folly. Sir Tuomas CHaLioner’s translation
(Elizabethan).

The publications already issued are:

1. Sir Tuomas Hobsy’s translation (Elizabethan) of Baldassare
Castiglione’s Courtier.

2. Mr. ASHBEE’S translation of the Treatises of Benvenuto Cellini
on Metal Work and Sculpture. 600 copies. A few still left. Price
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3. The Hymn of Bardaisan, the first Christian Poem, rendered into
English verse from the original Syriac, by F. CRAWFORD BuRKITT, of Trinity
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4. Bunyan’s Pilgrim’s Progress. Edited from the earlier editions by
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5. An Account of the Church of St. Mary Stratford Bow. By
OsBoRN HILLs, with an introductory chapter by C. R. ASHBEE. 250 copies.
All copies offered for sale have been disposed of. Price £1.

6. Shelley’s Adonais. Coloured wood-block frontispiece by C. R.
ASHBEE. Onvellum. Rubricated. 50 copies. [Out of Print.

7. The Poems of Shakespeare and Lyrics from his Plays,
in the orthography of the earliest editions. Edited by F. S. ELuis, with a
frontispiece by REGINALD SAVAGE, and a new alphabet of ‘ Bloomers’ designed
by C. R. AsHBEE. The book is small 4to. (9 in. by 7 in.), 254 pages, printed in
red and black, with limp vellum cover, edition limited to 450 copies.
Price £2 net. [Ozt of Print.

These volumes are published on behalf of the Essex House Press by
Mr. EDwarD ARNOLD, and can be ordered either from him or from any
Bookseller.

13

PICTURE BOOKS FOR CHRISTMAS
PRESENTS.

. Really and Truly; or, The Century for Babes.
By Mr. and Mrs. Ernest AMES. Twenty splendidly Coloured
Plates, with amusing verses, depicting the great events of the
nineteenth century. 4to., 3s. 6d.

Lama

2. Tails with a Twist. An Animal Picture-book,
inexpressibly funny. By E. T. RreEp, the famous Punch artist.
With verses by ‘ Belgian Hare.’ Twenty plates, each coloured in
one tint. 4to., 3s. 6d. Only a limited number of copies now
remain.

3. More Beasts for Worse Children. By H. B. and
B. T. B. Grotesque pictures in black and white, and inimitably
clever verses. 4to., with coloured cover, 3s. 6d.

4. Ruthless Rhymes for Heartless Homes. By
Col. D. STREAMER, with Pictures by ‘G. H.’ Oblong 4to.,
3s. 6d.

5. The Modern Traveller. By H. B. and B. T. B.
Fully Illustrated, with coloured cover. 4to., 3s. 6d.

‘The gaiety of nations has not received such a stimulus as this book ears for many a
merry Christmas.’— World.

6. A Moral Alphabet: In words of from one to seven
syllables. By H. B. and B. T. B. Fully Illustrated, with cover in
colour, 3s. 6d.

“H. B.’s uncommon humour is still quite unspoilt, and the pictures of B, T. B. are as
full of surprises and quaint touches as of old.’—/szs.

14
SOUVENIRS FROM THE HOLY LAND.

‘A souvenir of Christmas and its sacred and beautiful associations at
once peculiarly appropriate and attractive has been prepared by the
Rev. Harvey B. Greene, who has spent, it appears, three springs in
gathering and pressing, with the aid of native helpers, wild flowers from
the Holy Land. These have been tastefully and skilfully mounted in
books and on cards, which should please the eyes and touch the hearts
of many. The largest collection, containing seventeen specimens, bears
the title of “ Wild Flowers from Palestine”; a smaller set of twelve
examples is embraced in ‘‘ Pressed Flowers from the Holy Land”; while
the single Flower from the Christ Land is an ideal Christmas Card.
Boné-fides is guaranteed by printed letters from the British and American
Consuls at Jerusalem, Dean Hole of Rochester has written an introduc-
tion to the Flower Volumes, and Mr. Greene has himself supplied letter-
press descriptions and Scripture references. But the Syrian wild flowers
speak most eloquently for themselves.’— Scotsman.

Mr. Harvey Greene's deeply interesting souvenirs can be obtained
in the following forms :

1. WILD FLOWERS FROM PALESTINE.

Gatbered and Pressed in Palestine,
With an Introduction by the Very REV. S. REYNOLDS HOLE,

DEAN oF ROCHESTER.

Cloth elegant, 16mo., 4s. 6d.

2. PRESSED FLOWERS FROM THE HOLY
LAND.
Gatbered and Pressed in Palestine,
With an Introduction by DEAN HOLE.
Tastefully bound, 32mo., paper, 2s. 6d.

3. A FLOWER FROM THE CHRIST LAND.

A lovely Christmas Card containing a Single Pressed Flower.
Price 6a.

‘Consider the lilies of the field, how they grow.’

15
BIOGRAPHY AND REMINISCENCES.

Adderley. FRANCIS OF ASSISI. (See page 6.)

Alexander. RECOLLECTIONS OF A HIGHLAND SUBALTERN,
during the Campaigns of the 93rd Highlanders in India, under Colin Campbell,
Lord Clyde, in 1857-1859. By Lieutenant-Colonel W. GORDON ALEXANDER.
Illustrations and Maps. Demy 8vo., cloth, 16s.

Arnold. PASSAGES IN A WANDERING LIFE. By THomas
ARNOLD, M.A. Demy 8vo., with Portrait, 12s. 6d.

Boyle. THE RECOLLECTIONS OF THE DEAN OF SALISBURY.
By the Very Rev. G. D. BovLz, Dean of Salisbury. With Photogravure Portrait.
One vol., demy 8vo., cloth, 16s.

Clough. A MEMOIR OF ANNE J. CLOUGH, Principal of Newnham
ee amie: By her Niece, BLANCHE A. CLouGH. With Portraits.
vo., 12s. 6d.

De Vere. RECOLLECTIONS OF AUBREY DE VERE. Third Edition,
with Portrait. Demy 8vo., 16s.

Hare. MARIA EDGEWORTH: her Life and Letters. Edited by
Avcustus J. C. Harz, Author of ‘ The Story of Two Noble Lives,’ etc. With
Portraits. Two vols., crown 8vo., 16s. net.

Hervey. HUBERT HERVEY, STUDENT AND IMPERIALIST.
By the Right Hon. Eart Grey. Demy 8vo., Illustrated, 7s. 6d.

Hole. THE MEMORIES OF DEAN HOLE. By the Very Rev. S.
Reynoitps Hotz, Dean of Rochester. With Illustrations from Sketches by
Leech and Thackeray. Popular Edition. Crown 8vo., 6s.

Hole) MORE MEMORIES: Being Thoughts about England spoken in
America. By Dean HoLe. With Frontispiece. Demy 8vo., 16s.

Hole. ALITTLE TOURIN AMERICA. By Dean Hote. Illustrated.
Demy 8vo., 16s.

Hole. A LITTLE TOUR IN IRELAND. By ‘OxoniAn’ (Dean HOLE).
Illustrated by JOHN LexcH. Large crown 8vo., 6s.

Holmes. RECOLLECTIONS OF A LONDON POLICE OUR
By THomMas HoiMEs. (See page 3.)

Holland. LETTERS OF MARY SIBYLLA HOLLAND. Selected and
edited by her Son, BERNARD HOLLAND. Second Edition. Crown 8vo.,
7s. 6d. net.

Jowett. BENJAMIN JOWETT, MASTER OF BALLIOL. A Personal
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Cloth, 3s. 6d.

Le Fanu. SEVENTY YEARS OF IRISH LIFE. ‘By the late W. R.
Le Fanu. Popular Edition. Crown 8vo., 6s.

16

Macaulay. THE LIFE AND CORRESPONDENCE OF ZACHARY
MACAULAY. By Viscountess KNUTSFORD. (See page 5.)

Macdonald. THE MEMOIRS OF THE LATE SIR JOHN A.
MACDONALD, G.C.B., First Prime Minister of Canada. Edited by JosBPH
Pork, his Private Secretary. With Portraits. Two vols., demy 8vo., 32s.

Merivale.e THE AUTOBIOGRAPHY OF DEAN MERIVALE. With

Selections from his Correspondence. With Portrait, demy 8vo., 16s.

Morley. THE LIFE OF HENRY MORLEY, LL.D., Professor of
English Literature at University College, London. By the Rev. H. S.
SoL.y, M.A. With two Portraits. 8vo., 12s. 6d.

Mott. A MINGLED YARN. The Autobiography of EDWARD SPENCER
Mott (NATHANIEL GuBBINS). Author of ‘Cakes and Ale,’ etc. Large crown
8vo., 12s. 6d.

Pigou. PHASES OF MY LIFE. By the Very Rev. FRANCIS PIGOU,
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Rochefort. THE ADVENTURES OF MY LIFE. By HENRI ROCHE-

FORT. Second Edition. Two vols., large crown 8vo., 25s.

Roebuck. THE AUTOBIOGRAPHY AND LETTERS of the Right
Hon. JOHN ARTHUR ROEBUCK, Q.C., M.P. Edited by RopeRT EADON
LEADER. With two Portraits. Demy 8vo., 16s.

Simpson. MANY MEMORIES OF MANY PEOPLE. By Mrs. M. C.
SImPsoN (née Nassau Senior). Fourth Edition. Demy 8vo., 16s.

Stevenson. ROBERT LOUIS STEVENSON. By WALTER RALEIGH,
Professor of English Literature at University College, Liverpool. Second
Edition, Crown 8vo., cloth, 2s. 6d.

Tollemache. TALKS WITH MR. GLADSTONE. By the Hon. L. A.
TOLLEMACHE. With Portrait. Cloth, 6s.

Twining. RECOLLECTIONS OF LIFE AND WORK. Being the
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THEOLOGY.

Hole. ADDRESSES TO WORKING MEN from Pulpit and Platform.
By Dean HoLE. Crown 8vo., 6s.

Hole. FAITH WHICH WORKETH BY LOVE. A Sermon preached
after the funeral of the late Duchess of Teck. Vellum, Is. net.

Holland. ESSENTIALS IN RELIGION. Sermons preached in
Canterbury Cathedral. By Canon F, J. HoLLanp. Crown 8vo., 3s. 6d.

oye oes CHANGE IN RELIGION. By Onyx. Crown
vo., 38. 6d,

17

HISTORY.

Benson and Tatham. MEN OF MIGHT. Studies of Great Characters.
By A. C, Benson, M.A., and H. F. W. Tatuam, M.A., Assistant Masters at
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Fisher. FINLAND AND THE TSARS. By JosEPH R. FISHER, B.A.
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Gardner. FRIENDS OF THE OLDEN TIME. By ALICE GARDNER,
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trated, 2s. 6d, ,

Gardner. ROME: THE MIDDLE OF THE WORLD. By ALICE
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Milner. ENGLAND IN EGYPT. By Sir ALFRED MILNER, G.C.M.G.,
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Odysseus. TURKEY IN EUROPE. By OpyssEus. (See page 1.)

Oman. A HISTORY OF ENGLAND. By CHARLES OMAN, Deputy
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All Souls’ College, and Lecturer in History at New College, Oxford ; Author of
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Ransome. THE BATTLES OF FREDERICK THE GREAT. Extracted
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Rendel. NEWCASTLE-ON-TYNE: Its Municipal Origin and Growth,
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18

LITERATURE AND CRITICISM.

Bell KLEINES HAUSTHEATER. Fifteen Little Plays in German for
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Butler. SELECT ESSAYS OF SAINTE BEUVE. Chiefly bearing on
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Collingwood. THORSTEIN OF THE MERE: a Saga of the Northmen
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With Illustrations. Price 10s. 6d.

Cook. THE DEFENSE OF POESY, otherwise known as AN APOLOGY
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Cook, A DEFENCE OF POETRY. By Percy ByssHE SHELLEY.
Edited, with Notes and Introduction, by Professor A.S. Cook. Crown 8vo.,
cloth, 2s. 6d.

Davidson. A HANDBOOK TO DANTE. By GIovANNI A. SCARTAZZINI.
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M.A. Crown 8vo., cloth, 5s. :

Ellacombe. THE PLANT-LORE AND GARDEN-CRAFT OF
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Essex House Press Publications. (See pages 11 and 12.)

Fleming. THE ART OF READING AND SPEAKING. By the
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Garnett. SELECTIONS IN ENGLISH PROSE FROM ELIZABETH
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Goschen. THE CULTIVATION AND USE OF IMAGINATION.
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Harrison. STUDIES IN EARLY VICTORIAN LITERATURE. By
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Hudson. THE LIFE, ART AND CHARACTERS OF SHAKE-
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Kuhns. THE TREATMENT OF NATURE IN DANTE’S ‘DIVINA
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Lang. LAMB’S ADVENTURES OF ULYSSES. With an Introduction
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19

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Raleigh. STYLE. By WALTER RALEIGH, Professor of English Literature
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Raleigh. MILTON. By Prof. WALTER RALEIGH. (See page 5.)

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Rodd. THE CUSTOMS AND LORE OF MODERN GREECE,
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Schelling. BEN JONSON’S TIMBER. Edited by Professor F. E.
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POETRY.

Collins. A TREASURY OF MINOR BRITISH POETRY. Selected
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crown 8vo., 7s. 6d.

Crabbe. POEMS OF GEORGE CRABBE. Selected and Edited by
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Glencairn, R. J. POEMS AND SONGS OF DEGREES. By RoseErr J.
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Gummere. OLD ENGLISH BALLADS. Selected and Edited by
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Holland. VERSES. By Maup HoLLaND (Maud Walpole). Crown
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BY THE SAME AUTHOR.
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20

FICTION.

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‘ Adalet.. HADJIRA: A Turkish Love Story. By ‘ADALET.’ One vol.,
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Adderley. STEPHEN REMARX. The Story of a Venture in Ethics.
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Adderley. PAUL MERCER. A Tale of Repentance among Millions.
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Bunsen. A WINTER IN BERLIN. By MariE VON BUNSEN.
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Burneside. THE DELUSION OF DIANA. By MARGARET BURNESIDE.
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Charleton,. NETHERDYKE. By R. J. CHARLETON. One vol., crown

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Cherbuliez, THE TUTOR’S SECRET. (Le Secret du Précepteur.)
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Chester. A PLAIN WOMAN’S PART. By NORLEY CHESTER.
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Edwards. THE MERMAID OF INISH-UIG, By R. W. K. Epwarps.
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21

Falkner. MOONFLEET. By J. MEADE FALKNER. Second Edition,
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Ford. _ON THE THRESHOLD. By IsaBeLta O. Forp, Author of
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Gaunt. DAVE’S SWEETHEART. By Mary GAUNT. One vol., 8vo.,
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Hall. FISH TAILS AND SOME TRUE ONES. Crown 8vo., 6s.

Harrison. THE FOREST OF BOURG-MARIE. By S. FRANCES
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Hutchinson. THAT FIDDLER FELLOW. A Tale of St. Andrews. By
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Knutsford. THE MYSTERY OF THE RUE SOLY. Translated by
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Lighthall THE FALSE CHEVALIER. By W. D. LIGHTHALL,

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Spinner. A RELUCTANT EVANGELIST, and other Stories. By
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Wallace. LOTUS OR LAUREL? By HELEN WALLACE (Gordon Roy).
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Williams. THE BAYONET THAT CAME HOME. By N. WYNNE
WILLIAMS. Crown 8vo., 3s. 6d.

22

TRAVEL AND SPORT.

Bell. TANGWEERA: Life among Gentle Savages on the Mosquito Coast
of Central America. By C. N, BELL, With numerous illustrations by the Author.
Demy 8vo., 16s.

Beynon. WITH KELLY TO CHITRAL. By Lieutenant W. G. L.
Brynon, D.S.O., 3rd Ghoorkha Rifles, Staff Officer to Colonel Kelly with the
Relief Force. With Maps, Plans, and Illustrations. Second Edition. Demy
8vo., 7s. 6d.

Bottome. A SUNSHINE TRIP: GLIMPSES OF THE ORIENT.
Extracts from Letters written by MARGARET BOTTOME. With Portrait, elegantly
bound, 4s. 6d.

Bradley. HUNTING REMINISCENCES OF FRANK GILLARD
WITH THE BELVOIR HOUNDS, 1860-1896. Recorded and Illustrated
by CUTHBERT BRADLEY. 8vo., 15s.

Bull. THE CRUISE OF THE ‘ANTARCTIC’ TO THE SOUTH
POLAR REGIONS. By H. J. BuLL, a member of the Expedition. With
Frontispiece by W. L. WyLig, A.R.A., and numerous full-page Illustrations by
W. G. Burn-MurpocH. Demy 8vo., 15s.

Burton. TROPICS AND SNOWS: a Record of Sport and Adventure
in Many Lands. By Captain R. G. BuRTON, Indian Staff Corps. Illustrated,
demy 8vo., 16s.

Chapman. WILD NORWAY. By ABEL CHAPMAN, Author of ‘ Wild
Spain.’ With Illustrations by CHaRLES WHYMPER. Demy 8vo., 16s,

Freshfield. THE EXPLORATION OF THE CAUCASUS. By
Dovucias W. FRESHFIELD, F.R.G.S., lately President of the Alpine Club.
Illustrated with Photogravures and Maps, 2 vols., 4to., £3 3s. net.

Gleichen. WITH THE BRITISH MISSION TO MENELIK, 1897.
By Count GLEICHEN, Grenadier Guards, Intelligence Officer to the Mission.
Illustrated, demy 8vo., 16s.

Gordon. PERSIA REVISITED. With Remarks on H.I.M. Mozuffer-
ed-Din Shah, and the Present Situation in Persia (1896). By General Sir T. E.
Gorpon, K.C.I.E., C.B., C.S.I. Formerly Military Attaché and Oriental
Secretary to the British Legation at Teheran, Author of ‘The Roof of the
World,’ etc. Demy 8vo., with full-page Illustrations, tos, 6d.

Grey. IN MOORISH CAPTIVITY. An Account of the ‘ Tourmaline’
Expedition to Sus, 1897-98. By H. M. Grey. Illustrated, demy 8vo., 16s.

Hall. FISH TAILS AND SOME TRUE ONES. By BRapDNnock HALL,
Author of ‘Rough Mischance.’? With an original Etching by the Author, and
twelve full-page Illustrations by T, H. McLACHLAN, Crown 8vo., 6s.

23

Macdonald. SOLDIERING AND SURVEYING IN BRITISH EAST
AFRICA, By Major J. R. MACDONALD, R.E. Fully Illustrated. Demy 8vo., 16s.

McNab. ON VELDT AND FARM, IN CAPE COLONY, BECHUANA-
LAND, NATAL, AND THE TRANSVAAL. By Frances McNas. With
Map. Second Edition. Crown 8vo., 300 pages, 3s. 6d.

Pike. THROUGH THE SUB-ARCTIC FOREST. A Record of a
Canoe Journey for 4,000 miles, from Fort Wrangel to the Pelly Lakes, and down
the Yukon to the Behring Sea. By WARBURTON Pixg, Author of ‘The Barren
Grounds of Canada,’ With Illustrations by CHARLES WHYMPER, from Photo-
graphs taken by the Author, anda Map. Demy 8vo., 16s.

Pollok, FIFTY YEARS’ REMINISCENCES OF INDIA. By Lieut.-

Colonel PoLLox, Author of ‘Sport in Burmah.’ Illustrated by A. C. CORBOULD.
Demy 8vo., 16s.

Portal. THE BRITISH MISSION TO UGANDA. By the late Sir
GERALD PorTaL, K.C.M.G. Edited by Sir RENNEL Ropp, K.C.M.G. With
an Introduction by the Right Honourable Lord CromER, G.C.M.G. Illustrated
from Photos taken during the Expedition by Colonel Rhodes. Demy 8vo., 21s.

Portal. MY MISSION TO ABYSSINIA. By the late Sir Gerald H.
PorTAL, C.B. With Map and Illustrations. Demy 8vo., 15s.

Pritchett. PEN AND PENCIL SKETCHES OF SHIPPING AND
CRAFT ALL ROUND THE WORLD. By R. T. PritcHert. With 50
full-page Illustrations. Demy 8vo., 6s.

Reid. FROM PEKING TO PETERSBURG. A Journey of Fifty Days

in 1898. By ARNOT REID. With Portrait and Map. Large crown 8vo.,
7s. 6d.

Slatin and Wingate. FIRE AND SWORD IN THE SUDAN. By
Sir RupDoLF SLaTIN PasHa, K.C.M.G. Translated and Edited by Colonel Sir
F. R. Wincatsz, K.C.M.G. Fully Illustrated. Popular Edition. 6s. Alsoa
few copies of the Original Edition. Demy 8vo., 21s. net.

Smithh THROUGH UNKNOWN AFRICAN COUNTRIES. By A.
DONALDSON SMITH, M.D., F.R.G.S. With Illustrations by A. D. McCormick
and CHARLES WHYMPER. Super royal 8vo., One Guinea net.

Stone. IN AND BEYOND THE HIMALAYAS: A RECORD OF
SPORT AND TRAVEL. By S. J. Stons, late Deputy Inspector-General of
the Punjab Police. With 16 full-page Illustrations by CHARLES WHYMPER.
Demy 8vo., 16s.

Thompson. REMINISCENCES OF THE COURSE, THE CAMP,
AND THE CHASE. By Colonel R. F. Meysey THompson. Large crown
8vo., Ios. 6d.

Warkworth NOTES FROM A DIARY IN ASIATIC TURKEY.
By Ear Percy (then Lord Warkworth). With numerous Photogravures.
Feap. 4to., 21s. net.

24

THE SPORTSMAN’S LIBRARY.

Edited by the Right Hon. Sir HERBERT MAXWELL, Bart., M.P.

A Re-issue, in handsome volumes, of certain rare and entertaining books on
Sport, carefully selected by the Editor, and Illustrated by the best
Sporting Artists of the day, and with Reproductions of old Plates,

Library Edition, 15s. a Volume. Large-Paper Edition, limited to 200 copies,
Two Guineas a volume. Also obtainable in Sets only, in fine leather
bindings. Prices on application.

VOLUME I.

Smith, THE LIFE OF A FOX, AND THE DIARY OF A HUNTS-
MAN. By THomas SMITH, Master of the Hambledon and Pytchley Hounds.
With Illustrations by the Author, and Coloured Plates by G. H. JALLAND.

Sir RatpH Payne-Gatwey, Bart., writes: ‘It is excellent and beautifully produced.’

‘Is sure to appeal to everyone who has had, or is about to have, a chance of a run with the
hounds, and those to whom an unkindly fate denies this boon will enjoy it for the joyous music
of the hounds which it brings to relieve the winter of our discontent amid London fogs.’—PalZ

Mall Gazette.
‘It will be a classic of fox-hunting till the end of time.’— Yorkshire Post.
‘No hunting men should be without this book in their libraries.’ — Word.

VOLUME II.

Thornton. A SPORTING TOUR THROUGH THE NORTHERN
PARTS OF ENGLAND AND GREAT PART OF THE HIGHLANDS
OF SCOTLAND. By Colonel T. THorNTON, of Thornville Royal, in
Yorkshire. With the Original Illustrations by GARRARD, and other Illustrations
and Coloured Plates by G. E. Lopes.

‘Sportsmen of all descriptions will gladly welcome the pats new edition issued by Mr.
Edward Arnold of Colonel T. Thornton’s “ Sporting Tour,” which has long been a scarce book.’
—Daily News.

‘It is excellent reading for all interested in sport.'—Black and White.

‘A handsome volume, effectively illustrated with coloured plates by G. E. Lodge, and with
portraits and selections from the original illustrations, themselves characteristic of the art and
sport of the time.'— Times.

VOLUME III,

Cosmopolite. THE SPORTSMAN IN IRELAND. By a CoSsMOPOLITE.
With Coloured Plates and Black and White Drawings by P. CHENEVIX TRENCH,
and reproductions of the original Illustrations drawn by R. ALLEN, and engraved
by W. WeEsTALL, A.R.A.

‘ This is a most readable and entertaining book.'—Padi Mall Gazette.

‘ As to the “‘ get up” of the book we can only repeat what we said on the appearance of the
first of the set, that the series consists of the most tasteful and charming volumes at present
being issued by the English Press, and collectors of handsome books should find them not only
an ornament to their shelves, but also a sound investment.’

VOLUME IV.

Berkeley. REMINISCENCES OF A HUNTSMAN. By the Hon.
GRANTLEY F, BERKELEY. With a Coloured Frontispiece and the original
Illustrations by JoHN Lexcu, and several Coloured Plates and other Illustrations
by G. H. JALLAND.

‘The latest addition to the sumptuous ‘‘ Sportsman's Library” is here reproduced with all
possible aid from the printer and binder, with illustrations from the pencils of Leech and G. H.
Jalland.’—Globe.

‘The Hon. Grantley F. Berkeley had one great quality of the vaconteur. His self-revelations
and displays of vanity are delightful.’'— Times.

25

VOLUME V.

Scrope. THE ART OF DEERSTALKING. By WILLIAM SCROPE.
With Frontispiece by Epwin LANDSEER, and nine Photogravure Plates of the
original Illustrations.

‘With the fine illustrations by the Landseers and Scrope himself, this forms a most worthy
number of a splendid series.’—Pad] Mall Gazette.

‘Among the works published in connection with field sports in Scotland, none probably have
been more sought after than those of William Scrope, and although published more than fifty
years ago, they are still as fresh as ever, full of pleasant anecdote, and valuable for the many
practical hints which they convey to inexperienced sportsmen.’—Fredd.

VOLUME VI.

Nimrod. THE CHASE, THE TURF,AND THE ROAD. By NIMROD.
With a Photogravure Portrait of the Author by D. Mactisg, R.A., and with
Coloured Photogravure and other Plates from the original Illustrations by
ALKEN, and several reproductions of old Portraits.

‘Sir Herbert Maxwell has Ladle a real service for all who care for sport in republishing
Nimrod’s admirable papers. The book is admirably printed and produced both in the matter
of illustrations and of binding.’—St. ¥ames’s Gazette.

‘A thoroughly well got-up book.’—Wovid.

VOLUME VII.

Scrope. DAYS AND NIGHTS OF SALMON FISHING. By WILLIAM
Scrorg. With coloured Lithographic and Photogravure reproductions of the
original Plates.

© This great classic of sport has been reissued by Mr. Edward Arnold in charming form.’—
Literature.

COUNTRY HOUSE.

Brown. POULTRY-KEEPING AS AN INDUSTRY FOR FARMERS
AND COTTAGERS. By Epwarp Brown, F.L.S. Fully Illustrated by
Luptow. Revised Edition, demy 4to., cloth, 6s.

BY THE SAME AUTHOR.

PLEASURABLE POULTRY-KEEPING. Fully Illustrated. One vol.,
crown 8vo., cloth, 2s. 6d.

INDUSTRIAL POULTRY-KEEPING. (See page 3.)

POULTRY FATTENING. Fully Illustrated. New Edition. Crown 8vo.,
Is, 6d.

Cunningham. THE DRAUGHTS POCKET MANUAL. By J. G. CuN-

NINGHAM. An introduction to the Game in all its branches. Small 8vo., with
numerous diagrams, Is. 6d.

Elliot. AMATEUR CLUBS AND ACTORS. Edited by W. G. ELLIOT.
With numerous Illustrations by C. M. NewTon. Large 8vo., 15s.

26

Ellacombe. IN A GLOUCESTERSHIRE GARDEN. By the Rev.
H. N. Exvacomse, Vicar of Bitton, and Honorary Canon of Bristol, Author
of ‘Plant Lore and Garden Craft of Shakespeare.’ With new Illustrations by
Major E. B. RicKeTTs. Second Edition. Crown 8vo., cloth, 6s.

Gossip. THE CHESS POCKET MANUAL. By G. H. D. Gossip.
A Pocket Guide, with numerous Specimen Games and Illustrations. Small 8vo.,
Is. 6d.

Hole. A BOOK ABOUT ROSES. By the Very Rev. S. REYNOLDS
Ho ez, Dean of Rochester. Sixteenth Edition. Illustrated by H. G. Moon and
G. S. Etcoop, R.I. Presentation Edition, with Coloured Plates, 6s. Popular
Edition, 3s. 6d.

Hole. A BOOK ABOUT THE GARDEN AND THE GARDENER.
By Dean Hotz. Popular Edition, crown 8vo., 3s. 6d.

Holt. FANCY DRESSES DESCRIBED. By ARDERN HOLT. An
Alphabetical Dictionary of Fancy Costumes. With full accounts of the Dresses.
About 60 Illustrations by LiLL1an Younc. Many of them coloured. One vol.,
demy 8vo., 7s. 6d. net.

Holt. GENTLEMEN’S FANCY DRESS AND HOW TO CHOOSE
IT. By ARpDERN Hott. New and Revised Edition. With Illustrations.
Paper boards, 2s. 6d. ; cloth, 3s. 6d.

Maxwell. MEMORIES OF THE MONTHS. Second Series. By
- Sir HERBERT MAXWELL. (See page 2.)

“WYVERN’S’” COOKERY BOOKS.
Kenney-Herbert. COMMON-SENSE COOKERY: Based on Modern

English and Continental Principles Worked out in Detail. Large crown 8vo.,
over 500 pages. 7s. 6d.

BY THE SAME AUTHOR.

FIFTY BREAKFASTS : containing a great variety of New and Simple
Recipes for Breakfast Dishes. Small 8vo., 2s. 6d.

FIFTY DINNERS. Small 8vo., cloth, 2s. 6d.
FIFTY LUNCHES. Small 8vo., cloth, 2s. 6d.

Shorland. CYCLING FOR HEALTH AND PLEASURE. By
L. H. Porrer, Author of ‘ Wheels and Wheeling,’ etc. Revised and edited by

F. W. SHORLAND, Amateur Champion 1892-93-94. With numerous Illustrations,
small 8vo., 2s. 6d.

Smith, THE PRINCIPLES OF LANDED ESTATE MANAGE.
MENT. By Henry HERBERT SMITH, Fellow of the Institute of Surveyors ;
Agent to the Marquess of Lansdowne, K.G., the Earl of Crewe, Lord Methuen,
etc. With Plans and Illustrations. Demy 8vo., 16s.

White. PLEASURABLE BEE-KEEPING. By C. N. WHITE, Lecturer
to the County Councils of Huntingdon, Cambridgeshire, etc. Fully illustrated.
One vol., crown 8vo., cloth, 2s. 6d.

27

MISCELLANEOUS.

Bell. CONVERSATIONAL OPENINGS AND ENDINGS. By Mrs.
Hucu BELL. Square 8vo., 2s. 6d.

Clouston. THE CHIPPENDALE PERIOD IN ENGLISH FURNI-
TURE. By K. WarREN CLOUSTON. With 200 Illustrations by the Author.
Demy 4to., handsomely bound, One Guinea net.

Fell. BRITISH MERCHANT SEAMEN IN SAN FRANCISCO. By
the Rev. JAMES FELL. Crown 8vo., cloth, 3s. 6d.

GREAT PUBLIC SCHOOLS. Eton — Harrow — WINCHESTER —
RuGBy— WEsTMINSTER— MARLBOROUGH — CHELTENHAM — HAILEYBURY —
CLIFTON—CHARTERHOUSE, With nearly 100 Illustrations by the best artists,
Popular Edition. One vol., large imperial 16mo., handsomely bound, 3s. 6d,

Greene. FLOWERS FROM PALESTINE. (See page 14.)

HARROW SCHOOL. Edited by E. W. Howson and G. TowNsEND
WaRNER. With a Preface by EarL SPENCER, K.G., D.C.L., Chairman of the
Governors of Harrow School. And Contributions by Old Harrovians and Harrow
Masters. Illustrated with a large number of original full-page and other Pen-
and-ink Drawings by Mr. HERBERT MARSHALL. With several Photogravure
Portraits and reproductions of objects of interest. One vol., crown 4to., One
Guinea net. A Large-Paper Edition, limited to 150 copies, Three Guineas net.

Hartshorne. OLD ENGLISH GLASSES. An Account of Glass Drinking-
Vessels in England from Early Times to the end of the Eighteenth Century.
With Introductory Notices of Continental Glasses during the same period,
Original Documents, etc. Dedicated by special permission to Her Majesty the
Queen. By ALBERT HaRTSHORNE, Fellow of the Society of Antiquaries. Illus-
trated by nearly 70 full-page Tinted or Coloured Plates in the best style of Litho-
graphy, and several hundred outline Illustrations in the text. Super royal 4to.,
Three Guineas net.

Herschell. THE BEGGARS OF PARIS. Translated from the French
of M. Louis PauLian by Lapy HERSCHELL. Crown 8vo., Is.

Pilkington. IN AN ETON PLAYING FIELD. The Adventures of
some old Public School Boys in East London. By E. M. S. PILKINGTON.
Fcap. 8vo., handsomely bound, 2s. 6d.

ILLUSTRATED HUMOROUS BOOKS.
Ames. REALLY AND TRULY. By Mr. and Mrs. ERNEST AMEs. (See

page 13.)
H. B. and B. T. B. MORE BEASTS (FOR WORSE CHILDREN).
New Edition. One vol., 4to., 3s. 6d. (See page 13.)

BY THE SAME AUTHORS.
THE MODERN TRAVELLER. (See page 13.)

A MORAL ALPHABET. (See page 13.)

Lockwood. THE FRANK LOCKWOOD SKETCH-BOOK. Being a
Selection of Sketches by the late Sir Frank Lockwoop, Q.C., M.P. Third

Edition. Oblong royal 4to., ros. 6d.

28

Reed. TAILS WITH A TWIST. An Animal Picture-Book by E. T.
REED, Author of ‘ Pre-Historic Peeps,’ etc. With Verses by ‘A BELGIAN
Harg.’ Oblong demy 4to., 3s. 6d. (See page 13.)

Streamer. RUTHLESS RHYMES FOR HEARTLESS HOMES.
By Colonel D. STREAMER. (See page 13.)

SCIENCE AND PHILOSOPHY.

Arnold-Forster. ARMY LETTERS, 1897-98. By H. O. ARNOLD-
Forster, M.P. Crown 8vo., 3s. 6d.

Burgess. POLITICAL SCIENCE AND COMPARATIVE CONSTI-
TUTIONAL LAW. By Joun W. Burcsss, Ph.D., LL.D., Dean of the Uni-
versity Faculty of Political Science in Columbia College, U.S.A. In two vols.,
demy 8vo., cloth, 21s,

Graham. ENGLISH POLITICAL PHILOSOPHY: an Exposition and
Criticism of the Systems of Hobbes, Locke, Burke, Bentham, Mill and Maine,
By WILLIAM GrauaM, M.A., Professor of Jurisprudence and Political Economy
at Queen’s College, Belfast. Octavo, Ios. 6d. net.

Hill. A MANUAL OF HUMAN PHYSIOLOGY. By LEonarpD HILL
M.B. Nearly 500 pages and 170 Illustrations. Crown 8vo., 6s.

Holland. SUGGESTIONS FOR A SCHEME OF OLD AGE PEN-
SIONS. By the Hon. LIONEL HOLLAND. Crown 8vo., Is. 6d.

Hopkins. THE RELIGIONS OF INDIA. By E. W. Hopkins, Ph.D.
(Leipzig), Professor of Sanskrit and Comparative Philology in Bryn Mawr
College. One vol., demy 8vo., 8s. 6d. net.

Hutchison. FOOD AND THE PRINCIPLES OF DIETETICS.
By Dr. ROBERT HUTCHISON. (See page 7.)

Ladd. LOTZE’S PHILOSOPHICAL OUTLINES. Dictated Portions
of the Latest Lectures (at Gottingen and Berlin) of Hermann Lotze. Translated
and edited by GzorGE T. Lapp, Professor of Philosophy in Yale College. About
180 pages in each volume. Crown 8vo., cloth, 3s. 6d.each. Vol. I. Metaphysics.
Vol. II. Philosophy of Religion. Vol. III. Practical Philosophy. Vol. IV.
Psychology. Vol. V. 4sthetics. Vol. VI. Logic.

Lehfeldt. A TEXT-BOOK OF PHYSICAL CHEMISTRY. By Dr. R.
A. LEHFELDT, Professor of Physics at the East London Technical College.
Crown 8vo., 7s. 6d.

Morgan. ANIMAL BEHAVIOUR. By C. LLoyp Morgan, F.R.S.,
Principal of University College, Bristol. (See page 2.)

BY THE SAME AUTHOR.
ast AND INSTINCT: A STUDY IN HEREDITY. Demy 8vo.,
16s,

THE Sane OF CONDUCT. Cheaper Edition. Large crown 8vo.,
3s. 6d.

PSYCHOLOGY FOR TEACHERS. With a Preface by Sir JosHUA
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Third Edition. One vol., crown 8vo., cloth, 3s. 6d.

Paget. WASTED RECORDS OF DISEASE. By CHaARLEs E. PAGET,

Lecturer on Public Health in Owens College, Medical Officer of Health for
Salford, etc. Crown 8vo., 2s. 6d.

29

Pearson. THE CHANCES OF DEATH, and other Studies in Evolution.
By Karu Pearson, F.R.S., Author of ‘The Ethic of Free Thought,’ etc.
2 vols., demy 8vo., Illustrated, 25s. net.

Perry. CALCULUS FOR ENGINEERS. By Professor JOHN PERRY,
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Shaw. A TEXT-BOOK OF NURSING FOR HOME AND HOSPITAL
USE. By C. Weexs Suaw. Revised and largely re-written by W. RADFORD,
House Surgeon at the Poplar Hospital, under the supervision of Sir Dycz Ducx-
woRTH, M.D., F.R.C.P. Fully Illustrated, crown 8vo., 3s. 6d.

Taylor. THE ALPHABET. By Isaac Taytor, M.A., LL.D., Canon
of York. New Edition, 2 vols., demy 8vo., 21s.

THE JOURNAL OF MORPHOLOGY. Edited by C. O. WuHITMaN,
Professor of Biology in Clark University, U.S.A. Three numbers in a volume of
100 to 150 large 4to. pages, with numerous plates. Single numbers, 17s. 6d. ;
subscription to the volume of three numbers, 45s. Vols. I. to XIV. can now be
obtained.

Van ’T. Hoff£ LECTURES ON THEORETICAL AND PHYSICAL
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Young. A GENERAL ASTRONOMY. By CuHarues A. Youn,
Professor of Astronomy in the College of New Jersey, Associate of the Royal
Astronomical Society, Author of ‘The Sun,’ etc. In one vol., §50 pages, with 250
Illustrations, and supplemented with the necessary tables. Royal 8vo., half
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Dymond. CHEMISTRY FOR AGRICULTURAL STUDENTS. By
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Halliday. STEAM BOILERS. By G. HALuiDay, late Demonstrator at
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Wilson. ELECTRICAL TRACTION. By ERNEST WILSON, M.LE.E.,
Professor of Electrical Engineering at King’s College, London. Illustrated.
Crown 8vo., 5s.

THE NATIONAL REVIEW.
Edited by L. J. MAXSE.
Price Half a Crown Monthly.

The ‘National Review’ is the leading Unionist and Conservative
Review in Great Britain. Since it passed into the control and editor-
ship of Mr. Leo Maxse, most of the leaders of the Unionist Party have
contributed to its pages, including the Marquis of Salisbury, Mr. Arthur
Balfour, Mr. J. Chamberlain, and Lord George Hamilton. The episodes
of the month, which give a masterly review of the important events of

30

the preceding month, form a valuable feature of the Review, which now
occupies a unique position among monthly periodicals.

PUBLICATIONS OF THE INDIA OFFICE AND OF THE GOVERNMENT OF INDIA.
Mr. Epwarp ARNOLD, having been appointed Publisher to the Secretary of State for India in
Council, has now on sale the above publications at 37 Bedford Street, Strand, and is prepared to
supply full information concerning them on application.
INDIAN GOVERNMENT MAPS.
Any of the Maps in this magnificent series can now be obtained at the shortest notice from
Mr. EpwarD ARNOLD, Publisher to the India Office,

The following Catalogues of Mr. Edward Arnold's Publications will be sent post free on application:
CATALOGUE OF WORKS OF GENERAL LITERATURE.

GENERAL CATALOGUE OF EDUCATIONAL WORKS, including the principal publications of
Messrs. Ginn and Company, Educational Publishers, of Boston and New York.

CATALOGUE OF WORKS FOR USE IN ELEMENTARY SCHCOLS.
ILLUSTRATED LIST OF BOOKS FOR PRESENTS AND PRIZES.

BOOKS FOR THE YOUNG.

SIX SHILLINGS EACH.
FIRE AND SWORD IN THE SUDAN. By Sir Rupo.pH Satin and Sir F. R.

WincaTE. (See page 23.)

MOONFLEET. By J. MEADE FALKNER. (See page 21.)

FIVE SHILLINGS EACH.
SNOW -SHOES AND SLEDGES. By Kirk Mouwrog. Fully illustrated.
Crown 8vo., cloth, ss.

RICK DALE. By Kirk Munrog. Fully illustrated. Crown 8vo., cloth, 5s.
THE FUR SEAL’S TOOTH. By Kirk Munrog. Fully illustrated. Crown

8vo., cloth, ss.

HOW DICK AND MOLLY SAW ENGLAND. By M. H. CORNWALL

LrecH. With numerous Illustrations. Foolscap 4to., 5s.

DR. GILBERT’S DAUGHTERS. By MarcareT HarrizT MATHEWS.

Illustrated by Curis. Hammonp. Crown 8vo., cloth, 5s.

ERIC THE ARCHER. By Maurice H. Hervey. With 8 full-page Illustrations.

Handsomely bound, crown 8vo., 5s.

THE REEF OF GOLD. By Maurice H. HERVEY. With numerous full-page
Illustrations, handsomely bound, gilt edges, ss.

BAREROCK ; or, The Island of Pearls’ By HENRY NAsH. With numerous

Illustrations by LANCELOT SPEED. Large crown 8vo., handsomely bound, gilt edges, ss.

WAGNER’S HEROES, By Constance Maup. Illustrated by H. GRANVILLE

FELL. Crown 8vo., 5s.

WAGNER’S HEROINES. By Constance Maup. Illustrated by W. T. Maun.
Crown 8vo. 5s.
THREE SHILLINGS AND SIXPENCE EACH.
TALES FROM HANS ANDERSEN. With nearly 40 Original Illustrations
by E. A. Lemann. Small 4to., handsomely bound in cloth, 3s. 6d.

THE SNOW QUEEN, and other Tales. By Hans CHRISTIAN ANDERSEN.
Beautifully illustrated by Miss E. A. Lemann. Small 4to., handsomely bound, 3s. 6d.

gt

HUNTERS THREE. By Tuomas W. Knox, Author of ‘The Boy Travellers,’

etc. With numerous Illustrations. Crown 8vo., cloth, 3s. 6d.

THE SECRET OF THE DESERT. By E. D. Fawcett. With numerous
full-page Illustrations. Crown 8vo., cloth, 3s. 6d.

JOEL: A BOY OF GALILEE. By ANNiz FELLOws JOHNSTON. With ten
full-page Illustrations. Crown 8vo., cloth, 3s. 6d.

THE MUSHROOM CAVE. By Evetyn Raymonp. With Illustrations.
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THE DOUBLE EMPEROR. By W. Larrp Ciowss, Author of ‘The Great
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SWALLOWED BY AN EARTHQUAKE. By E. D. Fawcstt. Illus-
trated. Crown 8vo., 3s. 6d.

HARTMANN THE ANARCHIST; or, The Doom of the Great City. By

“E. Douctas Fawcett. With sixteen full-page and numerous smaller Illustrations by F T.
Jane. Crown 8vo., cloth, 3s. 6d.

ANIMAL SKETCHES: a Popular Book of Natural History. By Professor C,
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sat th THe MIDDLE OF THE WORLD. By Aticz GarDNer. IIlustrated.
loth, 3s. 6d.

TWO SHILLINGS AND SIXPENCE.

FRIENDS OF THE OLDEN TIME. By Aticz GARDNER, Lecturer in
History at Newnham College, Cambridge. Second Edition. Illustrated. Square 8vo., 2s. 6d.

TWO SHILLINGS EACH.
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Royal 4to., 7s. 6d. and 5s.

PAGE

ABOUT - - - - 20
‘Adalet’ - = 20
Adams, E. Davenport 31
20

Alexander, W. Gordon 15

Adderley, Hon. and Rev. J. 6,

Ames, Ernest - - 13
A Moral Alphabet - 13
Andersen, Hans Christian 30
Arnold-Forster, H. O. 2, 28
Arnold, Thomas : - 15
Ashbee, C. R. - - - 12
Bagot, Mrs. J. : - 4
Bell, Mrs. Hugh - 18, 27
Bell, Napier - - - 22
Belloc, Hilaire - - 8
Benson, A. C. - - - «17
Berkeley, Hon. Grantley F.- 24
Beynon, W.G.L. - + 22
Bottome, Margaret - - 22
Boyle, Very Rev. G. D. - 15
Bradley, Cuthbert = - - 22
Brown, Edward - 3) 25
Bull, H. J. - - 22
Bunsen, Marie von - 20
Burgess, John W.- 28

Burneside, Margaret -

Burkitt, F. Crawford - - 12
Burton, Capt. R. G. - -
aT. :

Butler, 18
Chapman, Abel : - 22
Charleton, R. J. - - 20
Cherbuliez, Victor - - 20
Chester, Norley : 10, 20

’ Children’s Favourite Series - 31

Children’s Hour Series - 31
Cholmondeley, Mary - 9, 20
Clough, Blanche A. - 15
Clouston, J. Storer - - 9
Clouston, K. Warren - - 27
Clowes, W. Laird = - * 31
Coleridge, M. E. - - 26
Collingwood, W. G. - 18, 20
Collins, J. Churton’ - - 19
Cook, Prof. A. S. - - 18
Cosmopolite : : - 24
Cunningham, J.G. - - 25
Davidson, Thomas - - 318
De Vere, Aubrey : - 55
Dunmore, Earlof = - - 20
Dymond, T. S. - - 29
Edwards, R. W. K. - = 20
Ellacombe, H.N. - 78, 26

Elliot, W. G. - 25
Essex House Publications It, 12

Falkner, J. Meade -

* BF
Fawcett, E. D. : - 31
Fell, H. Granville - - 19
Fell, R Rev. J. - - - 27
Fenton, Mrs. - : - 6
Field, Mrs. E. M. - 31
Fisher, J. R. - - - 8
Fleming, Canon - - 18
Ford, Isabella O. : - 21
Freshfield, Douglas Ww. - 22
Frye, Alexis - - 31
Gardner, Alice : 7
Garnett, J. M. . - 18
Gaunt, Mary : : - 21

Fndex to Authors.

es

PAGE
Gleichen, Count : + 22
Glencairn, R. J. - - 19
Gordon, Sir T. E. + 22
Goschen, Rt. Hon. G. J. - 18
Gossip, G.H. D. - - 26
Graham, W. - - - 28
Great Public Schools - - 27
Greene, Harvey - - 14
Grey, Earl - - + 15
Grey, H. M. - - - 22
Gummere, F. B. : + 19

Haffkine, Dr. W. M. S 9

Hall, Bradnock - 21, 24
Halliday, G. - - - 29
Hare, Augustus J. C. + 15
Harrison, Frederic - - 18
Harrison, S. Frances - - 21
Harrow School - - 27
Hartshorne, Albert - - 27
Herschell, Lady - + 27
Hervey, M. H. : - 30
Hill, Leonard - - - 28
Hoff, Dr. J. H. Van ’T. + 10
Hofmeyr, A. - - - 4
Hole, Dean - + 15, 16, 26
Holland, Bernard - . 15,19
Holland, Hon. Lionel - 28
Holland, Maud - - 19
Holland, Rev. F. J. - - 16
Holmes, Thomas . - 3
Holt, Ardern - : - 26
Hopkinson, A.M. - + 31
Hopkins, E. W. : - 28
Hudson, H. N. : - 18
Hutchinson, Horace G. = 21

Hutchison, Robert - 7, 28

Indian Office Publications -
Johnston, Annie Fellows + 31

Journal of Morphology - 29
Jowett, Benjamins - - 15
Becsineysricrere - - 26
Knox, T. W. - - 31
Knutsford, Lady - 5, 21
Kuhns, L. Oscar - - 18
Ladd, G.T. - . - 28
Lang, Andrew - - - 38
Le Fanu, W. R. * 15
Legh, M.H. Cornwall 30, 32
Lehfeldt, Dr. R. A. - - 28
Lighthall, Ww. D. - - 21
Local Series - - 31
Lockwood, Sir Frank - - - 29
Macdonald, Lt.-Col. J. R. 13, 23
Macdonald, Sir John A. - 16
Mathews, Margaret H. - 30
Maud, Constance - Ig, 30
Maxse, L. J. - 29
Maxwell, Sir Herbert- 2, 26
McNab, Frances - - 23
McNulty, Edward - + 21
Merivale, J. A. : - 16
Milner, Sir Alfred — - - 47
Modern aravellet, The - 23
Montrésor,

More Beasts for Worse Children z I 7
Morgan, C. Lloyd - 2, 28, 31
Morley, Henry - - 16

Mott, E.S. - - - 16

Munroe, Kirk
Nasb, Henry -
National Review

Oseaden: Maud .

Paget, Charles E. -
Pasley, Sir T. S. -
Pearson, Karl - :
Perry, Prof, John -
Pickering, Sidney -

Pigou, Very Ree Francis

Pike, Warburton :
Pilkington, E. M. S.
Pinsent, Ellen F. -
Podmore, C. T. -
Pollok, Lieut.-Colonel
Portal, Sir Sea He
Price, LLR

Pritchett, R. T. -

Quiller Couch, A. T. -

Radford, C. H. -
Raleigh, Walter -
Ransome, Cyril -
Raymond, Evelyn -
Reed, E. T. -
Reid, Arnot - :
Rendel, Hon. Daphne
Reynolds, Rev. S. H.
Richmond, Rev. Wilfrid
Roberts, Morley -
Rochefort, Henri -
Rodd, Sir Rennel -

Roebuck, Rt. Hon. J. A.

Roy, Gordon -
Russell, W. Clark = -
Schelling, Prof. F. E.
Scrope, William -
Shaw, C. Weeks :
Shorland, F. W. -
Simpson, Mrs. M. C. -
Slatin Pasha, Sir Rudolf
Smith, A. Donaldson -
Smith, H. H. :
Smith; Thomas -
Spinner, Alice - -
Sportsman Library -
Stevenson, :
Stone, S. J. :
Streamer, Colonel D.-
Tatham, H. F.W. -
Taylor, Isaac -

tree

N
=

Thompson, Col. R. F. Meysey

Thornton, Col. T. -
Tollemache, Hon. L. A.
Turkey in Europe -
Twining, Louisa -
Wallace, Helen -
Warkworth, Lord -
White, C. N. - -
TS Gea
illiams ynne -
Wilson, Ernest -
Wingate, Sr F.R. -
Young, Charles A. -

wad gilt)