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servable facts, so that he might gain somewhat

the same impression that he would from see-
LOWER ORGANISMS ing the phenomena themselves. To aid in
INmy recent volume on the "Behavior of this, I characterized as typical such phenomena
the Lower Organisms" are set forth certain as were fairly representative of the behavior in
views which have called forth discussion and general, while phenomena that were rare or
criticism.' I n response to many questions unique, like the reaction to the electric current,
which indicate that there is a general interest I said were not typical and gave an inadequate
i n the subject, I wish here to reexamine some idea of the nature of behavior. This, while
of the matters raised. strictly true, has led to misunderstanding;
Objections have centered about my discus- certain critics have assumed that I considered
&on of certain general theories of behavior, such phenomena as of no importance from any
particularly of the tropism theory, and of cer- point of view. I believe that there is nothing
tain applications of the theory of selection. in my treatment that gives ground for this
Some of the criticisms are clearly just; others4 assumption. Special cases of this form of
seem to me to rest upon misunderstanding, criticism will come up in later paragraphs.
while still others show actual differences of 2. The most important characteristics of be-
opinion. To set in a clear light these different havior have always seemed to me those shown
categories is my present wish. in the biological interrelations of the physio-
The only question of importance is : How far logical processes : in the relations of behavior
is there a real difference of opinion, among to preserving the organism, to supplying the
workers familiar a t first hand with the phe- requirements for metabolism, and in general to
nomena, i n regard to (1) the actual, experi- keeping the other physiological processes in
mental facts of behavior, (2) the general and progress. These adaptive or regulatory char-
important laws or principles underlying these acteristics of behavior furnished the problem to
facts. Divergences due to different lines of the solution of which attention was in my
interest, different fields of investigation and book mainly directed. Therefore I character-
different understanding of terms merely ob- ized as " important," " significant," and the
Mcure the essential point and need to be cleared lilre, mainly those features of behavior which
away. To make clear the objects and meaning seemed to lead toward an understanding of its
of different investigators sometimes reveals regulatory character. Other investigators, not
harmonious diversity in place of conflict ;when having this problem in the center of interest,
this result is not reached, it a t least shows pre- have considered quite other matters as the im-
cisely where opposition lies, and suggests ex- portant ones. Thus Torrey ( I . c.) holds that
perimentation that shall turn opposition into we find the most important features of be-
agreement. havior in precisely those features that are not
I t will greatly facilitate the attainment regulatory. Judgments of importance are of
of these ends if I first set forth briefly certain course relative; inlportance for zuftat? is the
purposes and principles that guided me in the question. I take i t that the question in which
preparation of my account. Torrey is primarily interested is that regard-
1. My book was designed mainly as a topo- ing the nature of the immediate change which
graphic survey of the field of behavior in the occurs in living matter when an external stim-
lowest organisms. My primary purpose was ulus acts upon it.
to give the reader a clear idea of the ob- 3. We now come more directly to the con-
'See especially Torrey, in SCIENCE,September tent of the work. Years of investigation had
6, 1907; Loeb, The Journal o f Bxpcrima+ztalZool- convinced me that the complexity of the prob-
ogy, Vol. 4, No. 1, 1907, and PfEiiger's Archiu, lems of bchavior had been underestimated;
Bd. 115, 1906, p. 580; Parker, SCIENCE, October that even in the lower organisms we are com-
25, 1907. pelled to deal with an immense number of

complex and little-understood factors, rather behavior must realize that there was such
than with a few simple ones. justification, even though he may himself hold
I n my book I therefore attempted, by cita- to some other definition of the word tropism.
tion of precise experimental data, to show the The great movement toward extreme simpli-
great number of factors which play a part in fication in these matters has certainly been
determining behavior; to show that changes in generally identified with the tropism theory;
the internal system which makes up the organ- " reduction to simple tropisms " has been the

ism are of equal importance for this, with ideal. Doubtless not all investigators have
changes in the external system which makes up held the tropism theory to be so simple and of
the environment; and I set forth particularly B U C ~wide applicability, but it is true, that
those remarkable relations of dependence and there has been a general belief that such was
support between the acts of behavior and the the c a s e - belief not confined to the unin-
other physiological processes, that are com- formed, but shared by workers of high stand-
monly spoken of as adaptation. ing. Thus, Bohn, in his recent admirable re-
4. My positive contention that behavior in view' of this entire question, after setting
the lower organisms is complex, varied and forth in detail examples of the tropism theory,
variable, so that it is not easily predictable, led says, "It is evident that nothing is simpler
me to a criticism of theories which represented than this explanation," and again, "For more
such behavior as simple, uniform. over wide than ten years certain biologists have thus
fields, and unequivocally determined by single explained the actions of animals by tropisms.
external factors. I found a theory of this .. . This had become the necessary and suf-
character to be widely held; I met i t in opposi- ficient explanation of all cases. Whenever i t
tion a t every turn as my papers began more was observed that animals accumulated a t a
and more to present behavior as complex; and point, without even seeking to determine how
I found this view presented regularly under they reached that point, a tropism was made
the name of the " tropism theory." to intervene." Bohn makes these statements
This then was the reason for my attack on merely as a presentation of well-known facts,
the tropism theory. I criticized it, not as a and it would be easy to multiply quotations
mere statement of one of the factors that make from biologists of the first rank showing that
up the complex phenomena of behavior, but as this idea of tropisms was a general one.
a view of supposed extremely wide applica- I n view of certain passages in Professor
bility, which maintained the simplicity and Loeb7s recent paper,' a note of historical char-
uniformity of the behavior of the lower organ- acter is here required. My criticisms have
isms. I tried to show that there was no single been directed not against any person or school,
schema into which most of the behavior of the but against a prevalent view. I have never
lower organisms could be forced. How far it considered any single person as the sole author
was just to identify tho view criticized with or only proper expositor of the tropism theory,
the tropism theory we shall inquire in a mo- but have taken the theory as I found i t com-
ment; here i t is most important, if we wish monly presented in biological literature. I
to get a clear understanding of the grounds of have not, therefore, considered i t necessary to
apparent conflict, $0 grasp the fact that the accompany a statement of my results with an
simplicity, unifolrmity and general application exposition of Loeb7s work and views ; there are
of a single schema were the points against other authors whose work and friendship I
which my criticism was directed. value highly whose expressed views are more
Was this idea of the tropism theory suffi- directly in opposition with what I have
ciently general to justify a criticism of it on pointed out than are Loeb7s. Certainly many
that basis? The word tropism has been used authors besides Loeb have ventured on inde-
ir, many senses and the theory haa taken many * Jourfial of Eloperimefital Zoology, 4, l51-156.
forms, as we shall see later; but I believe that a " Les Tropismes, les Reflexes et lYIntelYigence,"'

any one who has followed the literature of L'Afide Psyckologique, T. 12, 1906, pp. 137-156.

pendent contributions to the tropism theory diversity of view. The theories held by cer-
and on expositions of it; if i t was an error tain investigators give no ground for consider-
to take these into consideration, I am guilty ing the tropism a simple, elementary phe-
of that error. Here we are seeking to discover nomenon, nor one of wide application to lower
whether there is divergence of view as to the organisms; they involve a highly developed
facts themselves, and nothing would be more sensory apparatus and a complex activity of
gratifying than to find in Professor Loeb an the nervous system. Against such theories my
ally instead of an opponent, in this question criticism was not directed. On the other
of the complexity of behavior. We may, if hand, there is a widely prevalent theory of
we desire, call the theory which I criticized the tropisms which if correct really justifies the
popular tropism theory. common view of the elementary simplicity of
Now, how did this popular tropism theory these phenomena. This is the "local action
simplify behavior? There is among investi- theory of tropisms," and it was against this
gators an extraordinary diversity of opinion as that my criticism was directed.
to what a tropism is. Some use the word as a I wish to emphasize this point, as it gives
mere name for certain observed fa&. I n the key to the entire discussion. I found the
reading, conversation and correspondence I fountain head of the commonly held belief in
have met the following definitions, each held the simplicity and uniformity of the behavior
by well-known investigators : (1)any reaction of lower organisms in the "local action
of a lower organism is a tropism; (2) any theory "-representing the stimulus as pro-
reaction to the chemical or physical agents of ducing its reaction in that part of the body on
the environment is a tropism; (3) any move- which it directly falls, so that the organism re-
ment toward or away from a source of stimula- acts as a bundle of independent parts rather
tion is a tropism; (4) a tropism is any reaction than as a unit. I therefore attacked this
in which the organism turns as directly as pos- theory, and no other, in the chapter of my
sible toward or away from the source of stimu- book which deals with this matter. I believe
lation; (5) a tropism is any turning produced I made i t perfectly clear that this was the
by stimulation; (6) a tropism is any reaction theory under criticisin; in the title of the
i n which orientation to a steadily acting ex- chapter the '' local action theory of tropism "'
ternal force is the main characteristic. It is specified; I defined precisely what I meant
seems clear that there is no tropism "theory" by it; all through the chapter I took pains to
in any of these views; they merely apply a specify it, and in my summing up I expressly
name to certain facts, leaving the nature of the my statement of the tropism theory on page 94
reaction to be determined by experiment, and of my original paper is erroneous. The essential
permitting different explanations in different point in my characterization of the theory on that
cases. I myself a t first used the term (or its page seems to be the statement that "the theory
equivalent " taxis ") in some such collective of tropisms says that certain definite things hap-
sense, till a paper from the laboratory of one pen in the change of position undergone by organ-
of the leading exponents of the tropism theory isms under the influence of stimuli; that the
organisms perform certain acts in certain ways."
set forth with some warmth that the phe-
I f this is the point which Loeb holds to ba er-
nomena I described had "nothing to do with roneous, my criticisms of course do not touch his.
the tropisms." views in the least. Many authors present the
Among those who use the word tropism in a tropism theory as a theory of how reactions occur,
precisely defined sense, implying a theory as and it was as such that I criticized it. If I have
to the nature of the reaction,' there is likewise anywhere wrongly classified Professor Loeb with
I am uncertain whether Professor Loeb, in his these, I regret it, and am delighted to discover
recent paper (Jourm. E@p. ZooZ., 4 ) , wishes to my mistake. Any one who holds a theory ( o r
range himself with those for whom thk word would be a theory?) of tropisms that says noth-
tropism implies nothing as to the nature of the ing as to how the reaction occurs will hardly find
reaction. In his note on page 156, he says that anything in my discussion to oppose his views.
named the local action theory. I believe there port of which Parker and: Torrey cite the
h no ground for misunderstanding the theory circus movements of animals when the sense
that I was criticizing, though this seems to organs of one side have been obstructed. A
have occurred in certain cases. theory of such importance, it is contended,
My discussion has been attacked from two should have been dealt with i n a general work
eides. I n one recent number of Scr~iica: on the behavior of the lower organisms.
Parkern takes the ground that the local action Further, the chapter criticizing a theory under
theory is not held, so that it was not worth the name of tropism gives the impression that
while to demolish it; while in another recent this other theory is also condemned, though
number, Torreyo expressly defends the local ac- argumehts against it are not advanced.
tion theory. These mutually destructive criti- To ,this criticism my book is justly open. I
cisms naturally relieve me of some embarrass- should have given an exposition of the theory
ment in replying to both. Torrey's elaborate i n question, with an attempt to estimate the
defense shows that the theory is still very part it plays in the behavior of the lower or-
much alive and I can, therefore, only greet with ganisms. This unpurposed omission was
pleasure Parker's ready support of my main partly due to the fact that the two groups of
contention, even though this takes the f orm of whose behavior I gave a detailed exposition-
Sairey Gapp's crushing retort, " Who denige: the Protozoa and Ccelenterata-have furnished
of it, Betsey 2 Who deniges of it?" Parker's practically none of the evidence cited by my
work has been mainly with more complex ani- critics; partly to my attempt to focus atten-
mals than those dealt with i n my book, and his tion upon the local action theory as of in-
interests have lain rather in the field of sense &itely greater importance than the other form
physiology than in the development of activity. of the theory. B u t even though I held that
I can, therefore, readily understand that he action in accordance with the complex form of
should find it inconceivable that such a view the theory plays little part in the behavior of
~houldbe held; he has doubtless not met i t in the lower animals, the phenomena and theory
opposition a t every turn, as have those working should have been set forth, and I regret that
with the lowest organisms, and as I now meet this was not done.
it i n Torrey's paper. I n my book I have given We may now return to the criticisms and
precise statements of the theory i n the form defense of the local action theory. Regarding
of quotations from authors of highest stand- the nature of my criticisms, one point must
ing. Bohn7 in his recent exposition stdds be emphasized-a point that has been much
others. None of the authors quoted has, SO misunderstood, though I believe I expressed
far as I am aware, repudiated the local action myself explicitly on the matter. I made no
theory. It would appear, therefore, that a attempt, and had no desire, to deny the exist-
statement of the relation of the observed facts ence of the factors on which the local action
to this theory was much needed. theory, or any other existing theory of tro-
Before turning to the arguments urged in pisms, was based. So far as local action is
support of the local action theory, another concerned, I emphasized i n my book such cases
criticism of my discussion, made or implied as were established, and gave a list of them on
by most of my critics, must be dealt with. page 306. The question which I tried to an-
This may be put as follows. Suppose that the swer in my discussion of tropisms could be put
simple, local action theory of tropisms is not thus: After some years of study of the be-
satisfactory. Nevertheless, there is another, havior of the lower organisms, what is your
less precise, less simple, theory of tropisms impression regarding the extent and impor-
which is of itself important; a theory i n &up- tance of the part played by tropisms? A well-
@ SCIENCE, October 25, 1907. known investigator, after one of the most
'SCIENCE,September 6, 1907 ("the response to thorough and detailed studies of the behavior
$timulation is local," p. 319, etc.). of a certain group of invertebrates that have
'Loc. cit. ever been made, in which he watched and ex-

perimented with the animals literally day and long time been made with energy and per-
night for long periods, remarked to me, in dis- sistence. Our knowledge has grown till we
cussing this matter, that he never saw any are in a position to estimate the results. I n
tropisms. Without going so far as this, my the main it appears that to most of the be-
answer to the question aslied above was that havior the orientation idea has little appli-
"the theory of tropisms does not go far in cability. To that immense province of
helping us to understand the behavior of the behavior comprised in the reactions to chem-
lower organisms." I did not deny the exist- icals of all sorts (including food reactions,
ence of the phenomena which the theory takes respiratory reactions, etc.) it has shown it-
into consideration, but it seemed to me that self quite inapplicable. The case is the same
there are so many other factors, playing such with the reactions to heat and cold. With
important parts, that the tropism factor is of regard to the reactions to solid bodies nearly
relatively small importance in a general con- the same may be said, though there ara, some
sideration of behavior. I n my first paper on special cases in which the idea of orientation
this subject8 I included the more complex is applicable. The reaction to the electric
forms of the tropisms in this judgment. The current furnishes a typical orientation. I n
remark of the investigator above mentioned some of the reactions to gravity and to water
illustrates the fact that there are certainly currents the orientation idea is helpful. Yet
other aspects of behavior so striking and im- the recent worli of Lyon and others shows
portant as to quite mask the existence of that even in these the movement in a cer-
tropisms. tain direction is an essential part of the reac-
Let us attempt a brief characterization of tions ; they are essentially compensatory move-
tropisms, their history, and the part they play ments, and t.he taking of a certain orienta-
in behavior. tion is by no means the only important point.
1. The essential point in the tropism, as I n certain reactions to light the orientation
originally applied by Loeb to reactions to light idea has been most helpful, yet in an im-
was, in a word, the idea that the organism in mense proportion of the reactions of organ-
going toward or away from the light is not isms to this agent i t does not show itself
trying to go somewhere or to reach something, the essential point. The orientation theory
but is merely taking a certain position or is of greatest service in such cases as the
orientation in the light. This recognition going of insects toward a bright light, yet
that the positior, is the essential point was a even here such work as that of Holmes on
great step in advance, and its application by Ranatra shows that the orientation is not the
Loeb to certain features of the behavior of only point; the approach to the light seems
animals was an achievement of the highest after all essential, since if a certain orienta-
importance. tion does not bring the animal to the light,
2. This idea of orientation having proved it learns by experience to take a different
ao helpful in the study of reactions to light, orientation which does have this effect. The
the next step was, very properly, to apply work of Cole, of Radl and others, shows that
i t to other features of behavior, to see if i t in the lower organisms we have the begin-
would not prove equally useful elsewhere. nings of reactions to objects perceived
The reactions to chemicals, heat and cold, visually; the animal is not merely oriented
contact with solids, electricity, light, gravity, by the strongest light, but goes toward such
etc., were all brought under this point of view; objects, whether bright or dark, as might be
attempts to show that the position is the said to be "of interest ""0 it at the given
essential point in each of these have for a
The expression " of interest " of course has
"Contributions to the Study of the Behavior some objective equivalent, but to try to use it
of the Lower Organisms," Carnegie Institution, mould be to substitute an unintelligible conjecture
Publication 16, pp. 89-107. for an expression which at least conveys an idea
MAY 1, 19081 SCIENCE 703

moment. The reactions to light are bound up will not yield these conclusions. Single ob-
with almost every possible aspect of behavior, servations are of course important, since they
and the orientation principle plays in them are the material from which the large mass
but a relatively small part. is made, but single observations taken by
3. I n Loeb's original theory nothing was themselves do not help much in taking off the
said as to the way in which the position of facies of a long series of investigations, which
~rientationis reached, and I take it that he is what I tried to do. My conclusion, like
does not now consider this matter as belong- all statistical conclusions, is nothing that will
ing to the theory proper. But on this matter enable one to predict for a given individual
a tempting idea presented itself, to the effect case; if it were, it would of course be of much
that the position of orientation was reached greater value than it is. No single observa-
in the simplest possible way-by a local reac- tion whatever is inconsistent with my general
tion of the part on which the stimulating conclusion.
agent impinges. This gave the "local action Thus, writers who have flown to the defense
theory" which made the tropism a thing of of the existence of tropisms will find them-
such extreme simplicity; it has been applied, eelves in no conflict with my stand on the
in detail or in general, to all sorts of reac- matter. It was only the prevalent opinion of
tions, by many authors. While it holds in a the wide generality and importance of the
measure for the effect of the continuous elec- phenomena that I called in question. To hear
tric current, I believe that it has been that the actual existence of the tropism was
demonstrated that in the main this idea was held to require defense came as a real surprise
not correct; that the element it deals with to me. I f put forward as merely one factor
plays little part in behavior, aside from the out of many, with its relative importance sub-
effects of electricity. With this we shall deal ject to discussion, I shall agree most cordially.
in later paragraphs. I n certain quarters there seems to be an im-
Attempts have been made to controvert my pression that observation of the direct turning
position on the tropism theory by the per- of an organism toward a source of stimulation
formance of crucial experiments or by the is in some way opposed to my views, and that
citation of specific observations. These are citation of specific cases of this will come to
clearly based on misunderstanding. I t is me as a painful surprise. Yet, of course, this
obvious that the relative importance of an is one of the commonest and most evident facts
admitted single factor in producing a set of of behavior, and is discussed in detail in my
complex phenomena can not be settled in this book (see, for example, pp. 306-308). Its
manner. Valuable judgment on such a ques- existence is required if the theory I suggested
tion can be based only on an extended study is correct. I pointed out that in consequence
of the phenomena. My own opinion derives of the three factors in behavior whose impor-
any worth it may have solely from the fact tance I emphasized, this direct turning toward
that I had worked for nine years on the a localized stimulus would occur; if it did not
behavior of a large number of organisms, occur, that would tend to disprove the theory.
attempting to make a careful analysis, with "Innumerable instances of this class of reac-
tions could be given; they include perhaps the
detailed studies of the different factors in-
greater number of the directed movements of
volved and the part played by each. My con-
organisms " (p. 301).
clusions are of the same character as are
Thus the direct turning of animals is not in
drawn from a large mass of statistical data. dispute. A matter that is of interest lies i n
They can be adequately controverted only by the answer to the question whether the turning
showing that the acalysis of this mass of is due to the simple local reaction of the region
.data, or of another equally large or larger, on which the stimulating agent impinges. My
of the outward facts. The animals go toward own contention was that this is rarely the
~isiblethings that serve for food, protection, etc. case. I f authors will state clearly whether

they conceive the turning to be due to such vitiates its claim to consideration "; that " the
simple local reaction, and give the evidence on interesting phenomena of galvanic stimulation
which this opinion is based, that will be a real are to be so lightly put aside," etc., emanate
contribution on a disputed point. from Torrey, not from myself. So far have
The reaction to the electric current, i n which I been from " neglecting" it, that I devoted in
the effect is local and the behavior is unco- my book more space to this reaction than to
ordinated and unadaptive, is the type and pat- any other. But the importance of the reaction
tern of the local action style of behavior. Its to electricity is of the same sort in the be-
importance is thus naturally emphasized by havior of lower as in that of higher animals;
Torrey, in his defense of that theory, as though of the utmost importance, no one
against my own contention that this reaction would consider the reaction of a muscle to
is not typical of the behavior of lower organ- electricity " an adequate type of the behavior
isms. The question will be cleared most of mankind.)' I believe that it was made plain
readily by noticing the different objects which in my book that this was the point which I
guided us in taking our stands. My own pur- was setting forth.
pose, in my topographical survey of behavior, Torrey takes up my account of the reac-
was to give the reader a correct idea of the tions of Euglena to light, and attempts to
facts-of what he would see if he examined shorn that it agrees with what would be ex-
the phenomena himself. I n doing this, one pected from the local action theory of
must inevitably come, I believe, to my con- tropisms. It is not possible to take up the
clusion that the " action . . under the elec- details of this matter here. But I may point
tric current is not typical of the behavior out the following: I n accordance with my
under other stimuli." I f the reader examined general practise, my account i n this case was
accurately the reaction to the electric current based, not on an attempt to explain an iso-
he would see certain phenomena-local action, latcd reaction by a preferred theory, but on
lack of unity and coordination, different parts a n extensive analytical investigation of the
of the body opposing each other, etc. The reactions of the organism, attempting to iso-
question is-Is this typical of the behavior? late experimentally the various elements of
I s this what would be seen if the reactions to which the behavior is made up. I n this in-
heat, light, gravity, chemicals, etc., were ex- vestigation I was not able to find experi-
amined in the same way? Certainly i t is not. mentally that element which the tropism
I f the reader should get the impression that theory calls for, while those I did find ac-
the extraordinary series of phenomena seen counted for tho entire behavior. I, therefore,
when an electric current is passed through a had no ground for asserting the existence
collection of infusoria is likewise what is seen of the tropism element. I do not see that
when they are subjected to other stimuli, his Torrey has adduced any additional ground for
idea of behavior in the lower organisms would such assertion; at best he has merely tried to
be a ridiculous caricature of the reality. show that interpretation along the line he
There appears to be no reason for concealing prefers is not inconsistent with the facts.
this fact, and I set i t forth as clearly as I One of my figures of the reaction (Fig. 93
could. in my book) Torrey thinks "perfectly in
On the other hand, Torrey holds that in the harmony with the tropic schema," he says:
reaction to the electric current we may have "it is hard for me to conceive how an or-
exhibited in a very direct way some of the ganism swimming of necessity in a spiral
fundamental changes that occur in living mat- course could react more definitely to a moder-
ter when subjected to the action of a stimulus ; ate directive stimulus than Euglena does
hence its great importance. Nothing that I here "; and he " can only wonder at my run-
have said militates against this opinion. The ning so boldly and far into the enemy's
statements and implications that I hold that camp." Surely this last remark does not
"the uniqueness of the electric stimulus ... mean that Dr. Torrey considers i t a reputable

scientific procedure to manufacture or alter reactions the organism swerves, in becoming

a figure claiming to represent the facts, in oriented, only toward a certain side x, never
order to make i t agree with a theory. My toward the opposite side y, just as in the
figures were made from as precise a study of reaction of Euglena to light. But in the re-
the facts as I could make, before I had at- action to the electric current the organism
tempted by analysis of all the facts to see may be caused by the local action to swerve
what they mean, so that the figures form directly toward the side y, and to become
part of the data for my later conclusions. In oriented in that way. Local action would
my preface I said that my ideal was "to cause swerving toward the side 31 in the reac-
present an account that would include the tions to light, gravity, etc., exactly as in the
facts required for a refutation of my own electric current, and the fact that this does
general views, if such refutation is possible," not occur seems to be a demonstration that
and I should not be without the gratification local action is not the explanation in these
of having fuxlled that ideal if Torrey should cases.
be adjudged to have made out his case. But I t will then be clear, I hope, that my
the reason why I held that the organism does analysis was based on a thorough considera-
not react as directly as possible is as follows: tion of the available experimental data, and
The oriented organism is swimming toward not on prejudice for or against any given
the light in a spiral course, thus swerving first theory. Torrey indeed admits, if I under-
to the right, then to the left (omitting from stand him, the existence of all the factom
consideration the movement in other planes). which I set forth, and the correctness of
Now the light is changed, so as to come, say, my analysis so far as it deals with positive
from the right, as in my Fig. 95. The most factors, but believes that there is an addi-
direct way in which the organism, swimming tional factor, in virtue of which Euglena may
in a spiral, could become oriented to the light turn directly toward a light. Thus the
would be by an increase in the swerving to behavior of Euglena is more complex, accord-
the right and a decrease in the swerving to ing to Torrey's view, than I represented it.
the left, and this is what the tropism theory There is no doubt but that increase of knowl-
would lead us to expect. But the fact is that edge tends to reveal increased complexity in
there is an increase in the swerving both to the behavior of the lower organisms; of thie
the left and to the right, the spiral becom- many recent examples could be given. My
ing a wider one; the increase to the right
own work has had decidedly this tendency,
being, however, greater than that to the left,
but, as in the present case, I tried to keep the
the organism becomes gradually pointed to
theory as simple as the facts would permit.
the right. The increased swerving to the left
But my experiments on Euglena, while not
is not accounted for by the tropism theory,
and is indeed squarely opposed to it, while i t revealing the power of direct turning, do not
disprove its existence. It has always seemed
is to be expected if the analysis I gave is
correct. as extraordinary to me as to any one else that
The point becomes quite clear when we the direct turning should not occur. There
compare this reaction with that to the eleo is little profit in discussing matters which
tric current, which with its undoubted local only experiment can settle. At the time my
action Torrey considers a typical tropism. work was done, I had no opportunity to study
Since Euglena itself has not been shown to the reactions of Euglena in the stage when
react to electricity, we can not make the com- it has no flagellum and moves by contractions.
parison here, but Torrey does not maintain Such a study is much needed, and i t may
his views for Euglena alone, and the facts in reveal the additional factor which Tormy
the reactions of ciliates to light, gravity and looks for. Many higher organisms show a
water currents are parallel to those in the power of direct localization, in connection
reactions of Euglena to light. I n all these with complex activities of other sorts; there
706 SCIENCE IN. 8. VOL. XXVII. NO. 696

is no antecedent improbability of this in tion were kept in mind. With this, a stim-
Euglena. ulus, as the cause of a reaction, is lilrewise
Related to this matter is Torrey's discus- clearly defhed; this definition I had already
sion of the question whether the organism is given on page 6.
or is not stimulated after i t is oriented, which Stimulation and reaction are evidently, as
leads him finally to the extraordinary conclu- thus used, correlative terms; if there is no
sion that I "insist on an interpretation of reaction, there is no stimulation. I have no
organic behavior by means of general changa desire to insist that these are the only pos-
in internal states that are psychical rather sible definitions; I merely wish to point out
than physical," and to a general condemna- that this was my explicitly declared usage.
tion of my analysis on this account. Most NOF if we apply these definitions, the wholo
of the points made in Dr. Torrey's interest- structure of difficulties raised by Torrey falls
ing paper I can appreciate, but at the dis- to the ground. From the definitions i t fol-
cussion which leads to this conclusion regard- lows that when the movements of an organism
ing psychic factors I must confess my are uniform, it is not stimulated. After the
astonishment. The conclusion is reached infusorian has become oriented to light, it does
only by the aid of the somewhat desperate not change its movements, but swims in the,
assertion that to say that an oriented organ- same way it did before; there is then no ex-
ism is subjected to no general stimulation ternal evidence that it is stimulated, and if
"is no more than saying i t then possesses no my purely empirical definition is accepted, it
feeling of discomfort." Had I made such a is not stimulated.
statement, I should have expected much just I f i t be maintained, as Torrey does, that
and severe criticism for "psychologizing"; the organism is nevertheless stimulated at
for " crude anthropomorphism." such a time, then evidently some internal con-
The root of the difficulty lies in a misunder- dition is talren as a criterion of stimulation.
standing of certain of my attempts to avoid This is precisely the criterion which Torrey
the use of indefinite terms not having a pro- incorrectly attributed to me, and on the
cise experimental meaning; it comes finally ground of which he charged me with making
to ti simple matter of definition. Experi- " feelings of discomfort " and other psychical
mentally, it has seemed to me that the study phenomena the basis of my analysis. I f there
of behavior reduces mainly to a study of two were any sound foundation for his argument,
things: (1)the causes of changes in behavior; I could retort that i t is his view that calls
(2) the nature of the changes themselves. for the psychical factors. But, of course,
Now these two things correspond nearly to there is no reason for dragging in psychic
what are commonly called stimuli and reac- factors at all; it is perfectly easy to suppose
tions, though the common usage is a little that the organism when oriented is in a dif-
less precise, not always representing experi- fering physico-chemical state, and this as-
mental concepts. I, therefore, adopted for ex- sumed state might be considered stimulation,
perimental discussions the word reaction as unless the empirical definition of stimulation
signifying a change in behauior; the word as correlative with reaction is preferred. To
stimulus as meaning the cause of a change in be unable to conceive a change in physio-
behavior, though so far as I could I used the logical state otherwise than as psychical
plain phrases in place of the two terms. would seem to unfit one completely for the
Unless some such defhitions are used there objective analysis of behavior; such changes
is no experimental method of telling whether demonstrably occur even in unicellular organ-
an organism is reacting or not; whether i t is isms.
stimulated or not. On page 283 of my book It is evident that the highly objectionable
I took the greatest pains to emphasize the propositions which Torrey deduces from my
fact that my discussion would not be intel- discussion, to the effect that there can not
ligible unless this meaning of the word reac- be " a constant stimulus that does not induce

a differential movement," or that I deny the become more effective-more regulatory 1''
" possibility of symmetrical stimulation for The existence of unadaptive reactions not
an oriented organism," depend simply on the coming under this theory mas recognized.
definition of stimulus and reaction as correla- " The organism is composed of matter that is
tive terms of purely experimental meaning. ~ubject to the usual laws of physics and
I certainly believe that many animals, after chemistry. External agents may of course
they have ('fixed" a source of light and are act on this matter directly, causing changes
swimming toward it, are in a different physi- i n movement that are not regulatory >' (page
ological state from before. I f we deibe 345). The origin of these unadaptivo reac-
stimulation somewhat indefinitely as meaning tions I did not discuss, because I had no light
any such changed physiological state, then we to throw on the matter. But I emphasized
may of course hold that they are then stimu- my conviction that the study of the laws of
lated. There seems to be no real difference of matter and energy furnish the main field for
opinion on this matter; but the method of investigation, as compared with questions of
formulation of course depends on the defini- selection. "Whatever the part assigned to
tion of the terms employed. natural selection, the superlative importance
A further point discussed by Torrey has to of these laws remains; they must continue the
do with the relation between selection and chief field for scientific investigation" (p.
adaptation. As an aid to understanding the 326). I might have said "the only field,"
existence of adaptations in behavior, I ac- since of course the study of selection is merely
cepted certain forms of the selection theorj. the study of how these laws work under cer-
Torrey emphasizes the existence of unadap- tain complicated conditions.
tive reactions, like that to the electric cur- Torrey evidently overlooked my explicit
rent; he points out that there is no ground statements of the object and limitations of
for supposing that selection has played a part the theory in question.
in their production. To this I a p e e fully. This discussion of theories of development
B u t since Torrey draws therefrom the con- tends to give the impression that these form
clusion that "the hypothesis advanced by the important part of my treatment of be-
Jennings is not sufficiently broad to encom- havior. It is, therefore, only just to point out
pass all the phenomena it is devised to ex- that this matter was a side issue from the
plain," it needs to be pointed out that my main purpose of my work, and was explicitly
view was not " devised to explain " such phe- put forward merely as a suggestion as t o
nomena. A theory of selection, while directed what may have occurred. The short chapter
primarily to the explanation of adaptiveness, on this subject begins as follows: "It is not
requires the existence of raw material from the primary purpose of the present work to
which selection may occur, and this raw ma- treat the problems of development, but rather
terial must of course be largely unadaptive, to give an analysis of behavior as we now
or there would be no ground for selection. find it. But the results of this analysis fur-
$election can never account for the existence nish a certain amount of evidence as to how
of that from which selection is made. This, development may have occurred; this it will
I believe, was made plain in my book. '(It is be well to set forth briefly." The book is
clear that natural selection can not account primarily a treatment of behavior as a branch
for the origin of anything; only that can be of experimental physiology.
selected which already exists" (p. 326). I But I believe it to be short-sighted and un-
stated explicitly that the hypothesis set forth fortunate for a physiologist to attempt to set
was a theory of regulation; my exposition of i n opposition physiological interpretation, on
the matter opens on page 315 as follows: the one hand, with so-called "historical" in-
"The question in which we are interested is terpretation, dealing with selection and evolu-
then the following: How can behavior de- tion, on the other. Selection is not something
velop? That is, how can it change so as to outside of physiological or physico-chemical

action; it is merely a characterization of cer- use of psychic concepts i n explaining ob-

tain ways in which such action occurs. Some jective phenomena was i n full swing, I con-
of the relations brought about i n such action sidered that battle as fought and won; I have,
are lasting, while others are fleeting; those therefore, ever since proceeded, without dis-
which last are said ta be "selected." The cussion or ado of any sort, on that basis.
study of selection is an examination of the Every one must recognize the tremendous
relative permanency of different physico- service done by Loeb in championing through
chemical and physiological relations, and it thick and thin the necessity for the use of
is eminently a matter for experimentation. objective, experimental factors in the analysis
Again, i n addition to the rapid processes of behavior. No convinced experimentalist,
occurring mainly in the lifetime of individ- knowing the previous history of the subject,
uals, there are slow processes requiring more can reread, as I have just done, Loeb's early
than a generation to produce evident effects; work on behavior without being filled with
various aspects of these we call "heredity," admiration for the clear-cut enunciation, de-
" evolution," " genetics," etc. These slow fense and application of the principles on
processes belong as much to physiology as do which valuable experimental work has rested
the rapid ones. The existing condition of since that time, and on which it must continue
living things is known to be largely a product to rest.
of these processes, so that to attempt to ex- Any differences of opinion between Loeb
clude them from consideration and to act as and myself are then matters of detail; they
if their effects did not exist, when we are concern merely the results of the application
trying to understand living things, is a most of the agreed principles of investigation. It
futile proceeding. These matters are coming has seemed to me that some of the experi-
rapidly under experimental study, so that at- mentalists have rested content with superjicial
tempts to exclude them from consideration in explanations; that they did not realize the
physiology, as "historical," can not endure complexity of the problems with which they
much longer. were dealing. This has been the history of
We now come to the matter which seems to most applications of experiment to biology;
underlie most of the criticisms of my discus- the more thorough the work, the deeper are
sion. Certain authors seem to identify the the problems seen to be.
"tropism theory" with the view that the be- Thus I have not hesitated to bring forth
havior of organisms is to be explained by facts tending to show the inadequacy of the
objective, experimentally determinable factors. physico-chemical factors thus far set forth,
They feel that an attack on the "tropism the- and doubtless some have suspected that this
ory" is an attack on this view; this comes was done with the concealed purpose of dis-
forth notably in the criticisms made by Loeb crediting the general adequacy of such factors.
and Torrey, and i t is evident in the attitude This is a complete mistake; I did not till
of some other writers. lately realize even the existence of such a
There is, so far as I can see, nothing i n the suspicion. Complete confidence in the experi-
facts and relations which I have brought out mental method removes anxiety as to the effect
that i n any way opposes the principle that of criticizing the details of its application.
behavior is to be explained by objective, ex- My objections are only to the adequacy of
perimentally determinable factors--or indeed particular factors; they are based on experi-
that bears in any way on the question. I have mental grounds, and the difficulties they raise
simply assumed throughout that it is to be are to be resolved only by experimental study.
explained in that way, and I do not see how There is a vast difference between holding
experimental investigations can proceed on that behavior is fundamentally explicable on
any other basis. Beginning my work in 1896, experimental grounds, and holding that we
when the movement led by Loeb against the have already so explained it.

I n a recent paper Loeb* has intimated that ferent, and it is this common fundamental
even if the behavior of the organisms under principle to which the common name calls
consideration were as complex as that of man, attention. How far we should avoid words
the same objective and experimental methods that have ever had any psychic connotation
must be used in analyzing it. To this I fully whatever is a matter on which there may be
agree, and the behavior of man is of course divergence of opinion; but it is most impor-
no more to be excepted from this treatment tant to realize that this is totally distinct from
than is that of any other organism. I n some the question whether the psychic connotation
recent writings one finds indications of a is of any use in objective experimental an-
curious dualism, as if the behavior of lower alysis. I f this distinction is lost sight of,
organisms were to be analyzed in the objective, a divergence i n practical details is taken for
experimental way, but the behavior of higher a conflict in fundamental principles, to the
animals and man were not. This takes most detriment of experimental science. That i t
often the form of objection to any comparison is inlpossible to avoid such words completely
between the objective features of the behavior is seen when we find in the writings of such
of higher and lower animals, or to the use of men as Loeb the frequent use of such terms
the same terms in speaking of them, with a as " associative memory." Of course i t is to
tendency toward accusations of vitalism or only the objective phenomena that Loeb re-
" psychologizing," against those making such fers; but this is precisely the case also with
comparisons. Such accusations evidently de- other experimentalists accused of similar prac-
pend on the premise that the behavior of tises l
higher animals is to be explained only by To sum up the discussion with the defenders
vitalism or by "psychologizing." When one of the tropism theory: We all stand on the
is tempted to accuse an opponent in such same foundation, and the differences of opin-
ways, it is worth while to first examine whether ion are in matters of detail. I n attempting
the tendency to read psychic or vital factors to demonstrate the complexity of the problems,
into the phenomena does not lie in the mind of behavior, I have focused attention on a
of the accuser, rather than i n that of the certain precise and narrow form of the tropism
accused. When one has consciously and con- theory which seemed to me to have gained
sistently taken the ground that the behavior undue prominencein order to show that such
of all organisms, including man, is to be an- narrow schemata are inadequate. I n so doing
alyzed in the objective, experimental way, and other forms of the theory, more flexible in
that there is no ground for expecting a failure character, and setting forth the tropism as
of this method at any point, there is less occa- but one factor out of many, have been thrown
sion for anxiety at the use of similar terms into the background; of this the supportem
for similar objective phenomena throughout of the theory have justly complairied. With
the series. my main contention that behavior in the
For example, the "method of trial and lower organisms is complex7 involving many
error" is as much an objective phenome- factors, so that no one schema gives an ade-
non, to be explained by experimentally de- quate account of it, there seems to be little
terminable factors, in the dog or man, as in disagreement. As to the value of the "local
the infusorian. The undoubted great differ- action" theory there is still divergence of
ences between the exemplifications of the opinion.
"method" a t the two extremes are mat- And now a word as to my own positive con-
ters for experimental analysis and demon- tributions to the analysis of the matter. It
stration, if the experimental method is not is obvious that conclusions of the " statistical ''
to fail. They do not necessarily show that character that I have attributed to my own,
the fundamental principle involved is dif- are, from their relative inapplicability to ape-
Vf2iiger7r Archiv, 1906, 115, p. 681. cific cases, of much less value than precise
710 SCIENCE [N. 8. VOL. XXVII. NO. 696

ehemico-physical or physiological ones. It is physical properties of the mineral the reader

also obvious that to demonstrate the com- is referred to the original article.' The an-
plexity and difficulty of a field of work ia not alysis of the mineral, made by W. C. Blasdale,
a n achievement to be compared in value with showed the following :
6he demonstration that this field is simple and -
easily explicable on a few known principles. A
I am under no illusion in regard
- to this. The SiO, 43.56 43.79 43.68 0.723
clear-cut, narrow tropism theory would be of TiO, 20.18 20.00 20.09 0.250
infinitely greater value for predicting and - BaO 36.34 36.31 36.33 0.237
controlling the behavior of animals than any- /%0.08 100.10
thing I have offered, if only i t were true.
I am sure I regret that I can make no attempt This yields the empirical formula BaTiSi,O,.
to put an equally simple schema i n place of From this Louderback concludes: " Benitoite
the one I criticized; if the phenomena of be- is then a very acid titano-silicate of barium
havior were of elemental simplicity, that and stands in a class by itself, both as regards
would certainly be much more convenient, acid silicates and titano-silicates."
though perhaps they would then be less inter- Upon reading the paper, immediately
esting. Many of the concepts used in my after its publication last year, I noticed that
analysis-" physiological states," " selection," there was a very striking similarity to be
I' trial and error," and the like-are collective observed between the composition of benitoite
ones, characterizing varied phenomena of a and beryl, for, if benitoite be interpreted as a
Sigh degree of complexity. They all require metasilicate. we have:
much further analysis; they are programs for I1 111
Beryl .................. Be3A1,(Si03),

future work, not final solutions of the prob- I1 I V

lems. My analysis was mainly an attempt to Benitoite .............. Ba,Ti, ( SiO,) ,.

lay out the field, to point out the principal This similarity in the chemical composition
phenomena with which we have to deal, and is sufficient to consider the two compounds as
to define some of the main problems. I f any isomorphous, for, although titanium with a
one attempts to explain all behavior on any valency of four replaces aluminium with one
one basis, to unlock all its secrets by any of three, the total valences in both compounds
catchword whatever, be i t "trial and error," are the same. There is, however, a difference
," selection," " tropisms " or what not, he lacks
of one with respect to the number of atoms.
a realization of the complexity of his field of A few examples of well-known isonlorphous
investigation. Like other complex fields, that
series will show that the above is not unusual.
.of behavior, even in lower organisms, must be I n the marcasite group we have:
.divided up; the various factors must be sub- I1 I1
jected to long and intense special investiga- Marcasite ................... FeS S

tion, with a realization that we have here I11 I1

Arsenopyrite ................ FeAsS

material for the work of many generations I11 I11

s f investigators. H. S. JENNINCS Lijllingite ................... FeAsAs.

Here the number of atoms is constant in all

November 26, 1907

three compounds, but the valences vary. The
albite-anorthite group furnishes another illus-
I 111I V

OF' THE MINERAL BENITOITE * " Benitoite, a New California Gem Mineral,"
INJuly, 1907, Professor Q. D. Louderback University of California Publications, Bulletin
~ublished an interesting paper on the new of the Department of Geology, Vol. 5, 149-153,
mineral benitoite. For a description of the 1907.